To paraphrase Larry Agenbroad, the former director of the Mammoth Site in Hot Springs, SD, Mammoth taxonomy is confused, and confusing. And it has been this way for a long while. Henry Fairfield Osborn, a giant of North American vertebrate paleontology dedicated decades of his life (and that of his assistants) to the production of a 1600-page, 2-volume, tome describing the Proboscidea, published posthumously in 1942. Through a specimen-by-specimen analysis, he described 16 species of North American mammoths across 3 genera. Since that time, North American mammoth species have undergone significant pruning, with most paleontologists recognizing 4-5 species across North America: M. meridionalis (Southern Mammoth), M. columbi (Columbian Mammoth), M. primigenius (Woolly Mammoth), and M. exilis (Channel Island Pygmy Mammoth). A fifth species, M. jeffersonii (Jeffersonian Mammoth) was considered an intermediate form showing characteristics of both Columbian and Woolly mammoths.
The story went something like this…Around 1.5 million years ago, the Southern Mammoth (M. meridionalis) emigrated to North America, settling along the west coast. Shortly after, the Eurasian Steppe Mammoth (M. trogontherii) joined its trunked brethren. Both were found in early deposits in the Anza Borrego Desert of southern California (and potentially the Great Plains and Florida). The Southern Mammoth died out or was swallowed up by the more successful Columbian forms, which radiated throughout most of North America. The BIG mammoths that fill western museums, like Archie at the University of Nebraska and the Angus Mammoth at Denver were initially considered to be too big to be run-of-the-mill Columbian mammoths and were anointed “Imperial” mammoths. Woollies migrated down the front of the continental ice sheets late in the game, during the Wisconsin glaciation sometime in the last 100 thousand years. Jeffersonian mammoths were the love-children of Woolly and Columbian mammoths. And the Island Pygmies were early Columbian mammoths that swam the channel or wandered across a land bridge.
This was a great story. It had action and explanation. And it was the framework that most museums used to explain their monstrous Mammuthus mounts (or miniscule mounts, in the case of the Pygmy Mammoth on display at the Santa Barbara Museum of Natural History). But two papers in the last 6 months have shown that the reality is actually much more complicated…and interesting.
The first paper was a study by Adrian Lister and Andrei Sher. There are few scientists who have seen as many mammoths as Lister (who literally wrote the book on the subject). In a project that spanned decades, Lister and Sher visited many North American collections housing early mammoths. From California to Florida and everywhere in between. They concluded that the earliest mammoths on the continent were, in fact, not M. meridionalis. Rather they were an odd assortment of poorly prepared/reconstructed material, individuals with heavily worn teeth, or simply Columbian mammoths from an early context. The clincher was that they had an excellent sample of Old World Southern Mammoths that didn’t overlap with any of the North American specimens. Any-of-them…
The second paper (Poinar et al. in press) came out this week. A few years ago, we hosted a sharp graduate student from McMaster University (Ontario) who was interested in our midwestern mammoths, Jake Enk. I bought him lunch. We talked at length about messed up mammoth taxonomy. Normal stuff. Ultimately, Jake sampled ~30 teeth for genetic studies (like this one), then moved on to major collections of Mammuthus in Nebraska, Denver, UC-Berkeley, and Santa Barbara. Given the success rate of previous aDNA studies, we expected that one or two of these specimens might actually give us some decent data. To my surprise, Jake was able to extract complete mitochondrial genomes from 67 mammoths from south of the Laurentide ice. An even bigger surprise, was that they were all chips off of the same block. They weren’t even different species.
Well…this was a surprise/not surprise. We had been looking at this issue through the morphology of midwestern mammoth teeth and found that there was a significant amount of overlap between different “species” and shared our data with Jake. The conventional wisdom that Columbian mammoth teeth were distinct from woolly and Jeffersonian mammoth teeth just wasn’t holding up. You could see multiple morphs within a small geographic area–and we had the dates to prove that we weren’t seeing the influx of “new populations” through time. Things seemed to get really complicated in ecotonal areas, like Iowa. During the Last Glacial Maximum Iowa was a transitional landscape between the more open steppic grasslands to the west (“Columbian” mammoth territory) and the forest steppe (think Taiga) of the east (“Jeffersonian” and “Woolly” Mammoth territory). We hit collections at the University of Iowa, Iowa State Historical Society, Putnam Museum (Davenport) and the Sanford Museum (Cherokee) hard, hoping to figure out where one species left off and the other began. We ended up scratching our heads over animals that had jaws that looked like Woolly mammoths, but teeth that were Columbian…or jaws that were Jeffersonian on one side, but Woolly on the other. We found localities like the mammoth bonebed in Mahaska County, that had one jaw that looked like a Columbian mammoth, but two more that were dead ringers for big Woollies. These were exactly the morphological patterns that we might expect if a) Mammoths were a single biological population capable of inter-breeding and producing viable offspring, and 2) the midwestern mammoths were in the middle of the mess, showing characters of both populations.
So do these different “species” of mammoths mean anything? Why bother measuring teeth if all mammoths are the same? After the initial shock wore off I had plenty of time to think about this. As a morphologist, the idea that we are dealing with a single, morphologically variable population is actually…well…kind of liberating. Now we can explore how certain characters may have been selected for in different environments. We can think about functional morphology, broad-scale impacts of landscape/diet on body-size, or the morphological effects of introgressing populations. Before…the pygmy mammoths of California’s Channel Islands were an unrelated off-shoot of my midwestern behemouths, perhaps responding to nutritional stress and landscape changes in very different ways than their mainland cousins. Now, they are just another mammoth population that is using the same set of morphological and genetic tools to deal with the situation at hand. And we can learn from that.
Mammoths are fun to think about, even when we don’t know all of the answers. These papers (and a third that Jeff Saunders and I are hoping to finish up this week) illustrate the importance of retaining natural history collections in museums. A decade ago, there would have been no chance of getting this degree of genetic recovery out of fossil mammoths south of the ice. Even for “traditional” studies of morphology the only way to get sample sizes large enough to say meaningful things about the biogeography of a creature is to rely on the materials collected and accumulated through many generations.
Note: This is the first post in a series focused on a 4-year, National Science Foundation funded project to look at the extinction of Mammoths and Mastodonts in the Midwest.
For the last few years we’ve been traveling…a lot. We started a project in 2011 to better understand 1) when mammoths and mastodonts went extinct, and 2) the ecological mechanisms that might have played a major role in how they went extinct. The major foundation of this project is a museum-by-museum survey of mammoths and mastodonts in collections from nine states and one province (MN, WI, IA, MO, IL, IN, OH, KY, MI, and ON). Over the last 2.5 years, we’ve documented mammoths and mastodonts from 576 localities.
When we started this project, we knew that the Midwest was a hotbed for Pleistocene proboscideans. A compilation of known/published localities showed a continent-wide distribution, but definitely a concentration in the Great Lakes. Of course, as with most things paleontological, the best represented individuals are the youngest, and both genera overlap with the first humans in to the New World. The last standing mammoths on the continent are widely separated in space, found from the South Dakota Badlands to upstate New York. After the last-glacial maximum, mastodonts seem to be limited to forested areas of the Great Lakes region and Northwestern North America.
Mammoth and mastodont studies lie at the intersection of major research questions in a number of different disciplines. The reason that they are so important is primarily due to the fact that they are so common and widespread in the fossil record. Why, you ask? Probably size and distribution. Their remains are big enough to be seen from the cab of a tractor or backhoe, and were distributed coast to coast during the last half of the Pleistocene. Since they are relatively widespread and common components of the fossil record, we can get an elephant’s eye view of ecological changes, IF we know what questions to ask. Their remains are also much more common in museum collections than other victims of the terminal Pleistocene extinction event, so they might give us a glimpse into HOW the extinction occurred.
Why (Part I): Preposterous Proboscidean Paradigm Shifts
The 2005 discovery of a mammoth tusk in the bed of Sugar Creek (central Illinois) started it all. Dennis Campbell, biology professor at Lincoln College (and ISM research associate), had brought a class out to the creek to census freshwater mussels when Judd McCullum, (then a student in the class), stumbled across a large cylindrical object. Despite good-natured ribbing that it was “just a tree trunk”, Judd was convinced it was a mammoth tusk…and he was right. ISM paleontologist Jeff Saunders identified the tusk as a woolly mammoth. Conventional thinking had woolly mammoths in Illinois at the same time as the glaciers. We thought that they occupied the narrow band of tundra in front of the massive continent-grinding glaciers that covered the Midwest up until ~18,000 years ago.
To be thorough, Jeff submitted a sample for radiocarbon dating anyway. The results were surprising. Rather than dating to the time of the glaciers, the Lincoln College mammoth dated a few thousand years later, when central Illinois was covered by a cold swamp, with black ash and spruce as the dominant vegetation, not a grassland. This was a game-changer. Not only were woolly mammoths found outside of their traditional tundra habitat, but when the glaciers left the area, they stayed and survived in changing Midwestern ecosystems until their extinction, ~12,000 years ago.
Meanwhile, a graduate student at the University of Utah developed an interest in the ancient DNA of North American mammoths. Jake Enk, now finishing his PhD at MacMaster University in Ontario, managed to extract a good chunk of mitochondrial DNA from the Huntington Mammoth in Utah. The Huntington mammoth is the epitome of a Columbian Mammoth. It’s from the heart of the Columbian mammoth range, Utah. It’s cheek teeth, although fairly worn (this animal was 55-60 years old), consist of 7+ enamel ridge-plates spread out into a relatively long tooth (~6 plates per 10 cm). For good measure, Enk also extracted DNA from two additional Columbian mammoth teeth from Wyoming. Surprisingly, when compared to woolly mammoth DNA from Alaska and Siberia, these Columbian mammoths were similar. Actually, they were VERY similar. The three Columbian mammoth mtDNA sequences nested nicely within one of the Alaskan woolly clades. The take home message was that morphological variability in mammoths is much greater than genetic differences. These were not separate species–they probably don’t even merit being a sub-species.
But mastodonts were not immune to paradigm shifts. In 2011, Neal Woodman and Nancy Beaven published a report on the dating of a mastodon in northern Indiana, the Overmyer mastodon. The date they reported was 1500 years younger than expected. The typical pattern was that mastodonts went extinct ~12,900 BP, only a few hundred years after the first major human cultural group (Clovis) appears on the scene. The Overmyer animal, if the dates were to be believed, meant that mastodons survived not only the first wave of human colonization, but lived side-by-side with human groups almost into the Holocene!
Studies like these got us to thinking. What if there are other assumptions about the habitat preferences and behaviors of mammoths and mastodonts that we are wrong about? What would happen if we dated more specimens–or used new techniques for insight into paleodiets and behavior (i.e., stable isotopes) or population dynamics (i.e., ancient DNA)? Was the Lincoln College Mammoth the exception? Or the rule? What do the major morphological differences between different mammoth populations mean if they don’t reflect relatedness–or evolutionary history? Did Mastodonts really hang on so late? Why was there such a large gap between the Overmyer mastodont and other dated animals in the Midwest? All of a sudden, there were a lot of questions that we didn’t know the answer to.
Why (Part II): Elaborate Extinction Scenarios needing Evaluation!
These questions are important not only for understanding past ecological conditions, but for understanding one of those BIG questions…why did 35 genera of North American megafauna (species >100 kg) go extinct at the end of the last Ice Age? This extinction event is considered one of the BIG 5 mass extinctions in the history of life on Earth. Yet it is unique from earlier mass extinctions. In addition to being the most recent, the majority of the victims were the largest of the large fauna. Small fauna were spared, more or less, or managed to migrate to new ranges. Furthermore, the extinction of these species coincided with major climate changes AND the introduction of a novel, supposedly predatory species known to profoundly alter its environment and potentially overhunt its prey, Homo sapiens. The discussion surrounding this extinction event in recent years has become increasingly polarized. There are a number of scenarios that have been proposed to explain this extinction. Perhaps some of the megafauna were killed off by colonizing human populations, with the rest doomed as the result of ecosystem reorganization after the loss of keystone species such as mammoths. Alternatively, abrupt climate changes may have stressed megafaunal populations to the breaking point. Deglaciation was not simply a gradual warming. The glacial spring came in stops and starts, and may have presented megafaunal populations with a moving target. Never quite able to adjust to changing conditions. These are the main working hypotheses, but of course, there are others. Was it the mid-air explosion of a comet over glacial ice in Canada? A hypervirulent disease? A combination of the above? It is hard to say without more hard data on the timing and ecology of key extinct species such as mammoths and mastodonts. Beware of TV documentaries claiming that we now know the answer to what caused these extinctions. Most scientists agree (although there a vocal few who don’t) that we don’t have enough data to tease out the smoking gun…let alone identify who or what pulled the trigger!
How (Part I): New Techniques
But how do you tackle something as big as megafaunal extinctions? This is a global pattern involving many different species and ecosystems. What sort of data do you need to distinguish between different extinction scenarios? Obviously, timing is everything. In the last decade or so, direct dating of megafaunal bones has become more accurate and commonplace. For this project, we’ve been dating a lot fossils from museums, trying to fill in the gaps in space and time. We hope to say something about when these animals ultimately went extinct using new and improved chronological datasets. We also believe that animal ecology is an important aspect of survival, so we are utilizing techniques that capture the details of individual life histories. Specifically, chemical signatures from bones and teeth (in the form of stable isotopes) that can tell us about animal diets and mobility. (more on what we are learning from these techniques in future posts)
How (Part II): The Team
Modern paleontology does not happen without a team of experts, each providing critical data for hypothesis testing. This project is a collaboration between many different experts. Jeff Saunders (ISM) and myself are vertebrate paleontologists/paleoecologists who are tasked with understanding biogeographic variation in space. Stacey Lengyel (ISM) is an expert in dating techniques–she also happens to be creating a great website on Ice Ace mammals that will be launched this spring. Greg Hodgins is a bone chemist and dating expert at the University of Arizona. J. Douglas Walker (University of Kansas) and Alan Walker (Iowa State University) are experts is different types of isotopic analyses. Others have also contributed to our understanding of proboscidean paleoecology. Veterinarian Dennis Lawler (ISM) has been instrumental in exploring the impact of disease on mammoths and mastodonts and Eric Grimm (ISM) has provided environmental context for dated specimens through his work on ancient pollen recovered from the mud of midwestern lakes.
As we scale back the data acquisition phase of this project and focus more on analyzing the datasets that we’ve collected, we’ll have more to say about how mammoths and mastodonts lived and died, at least across the Midwest. A significant component of this project is dedicated to communicating our results to the public, primarily through online resources like this blog and the aforementioned website. So stay tuned for future developments. The data have started rolling in.
Enk, J., Devault, A., Debruyne, R., King, C. E., Treangen, T., O’Rourke, D., … & Poinar, H. (2011). Complete Columbian mammoth mitogenome suggests interbreeding with woolly mammoths. Genome biology, 12(5), R51.
Saunders, J. J., Grimm, E. C., Widga, C. C., Campbell, G. D., Curry, B. B., Grimley, D. A., … & Treworgy, J. D. (2010). Paradigms and proboscideans in the southern Great Lakes region, USA. Quaternary International, 217(1), 175-187.
Woodman, N., & Beavan Athfield, N. (2009). Post-Clovis survival of American mastodon in the southern Great Lakes region of North America. Quaternary Research, 72(3), 359-363.