Researching Fossil Ungulate Communities

alces-alces-porkkala-finland

Alces alces (moose), Porkkala, Finland; photo courtesy of Juha Saarinen

In their paper “Patterns of diet and body mass of large ungulates from the Pleistocene of Western Europe, and their relation to vegetation,” published this past September in Palaeontologia Electronica, Juha Saarinen, Jussi Eronen, Mikael Fortelius, Heikki Seppä, and Adrian Lister investigate fossil ungulate communities found in England, Ireland, and Germany.

Not fossil ungulates, fossil ungulate communities.

The variety of fossils studied is just one of the exciting elements of their research.  Rather than focusing on a single species—which, given the limitations of the fossil record, is usually the case—they studied groups of fossils from at least 14 different ungulate species from the Middle to Late Pleistocene.

“[W]e are now at a point,” wrote Juha Saarinen, lead author of the paper, in an email, “where enough fossil material of ungulates and pollen records have accumulated to enable such a large scale quantitative comparison of body size and diets of ungulate with local vegetation patterns in the past as we did. Comparing vegetation proxies and mammal ecometrics from fossil data using such quantitative statistical analyses as we did has, to our knowledge, never been attempted before, so that is probably the most novel achievement of this study.”

The ungainly name of ‘ungulate’ refers to hooved animals: even-toed and odd-toed (Artiodactyla and Perissodactyla, respectively). Examples include horses, deer, moose, rhinoceros, bison, pigs and hippopotamuses.

beneski-brontops-tyleri-brontothere-view-from-skull

Brontops tyleri (a type of brontothere and a Perissodactyl) at the Beneski Museum at Amherst College, Massachusetts.   Brontotheres survived until the Eocene, an era that ended approximately 30+ million years BEFORE the Pleistocene, so this animal–although an ungulate–was not part of this study. Picture taken by the author of this blog

 

Using mesowear on the fossil teeth, they were able to determine information about their diets (from browsing to grazing), and by comparing this data with the pollen record associated with the areas in which these fossils were found, they were able to tell whether they ate more browse or grass in either open or closed environments. Body mass for these fossils was calculated and then compared to the diet of these animals.

They were searching for answers to how these species adapted to the environment in which they lived.  How did their body size relate to the vegetation available? Was their body size influenced by possible predators or by other members of their species? (In other words, were they bigger to intimidate predators or were they smaller because they lived in expansive herds?) Or was thermoregulation the single determining factor in how big these animals became, as has been proposed in earlier studies?

 

Beneski - Irish elk

Megaloceros giganteus (otherwise known as Irish Elk and an Artiodactyl) in between a mastodon and a mammoth fossil at the Beneski Museum at Amherst College, Massachusetts; picture taken by the author of this blog

 

It interested me to learn that they relied on what I rather simplistically referred to as the ‘physical observation’ of fossils.

Mesowear analysis looks at the wear and shape of fossil teeth.  Various plant material affects tooth-wear in distinctly different ways, which can be seen both on the teeth themselves and in the way the teeth have evolved.

To be clear, “this is specifically wear-induced shape, not the original shape of the unworn teeth,” Juha added. “In other words, mesowear is the change in the shape of the teeth as they get worn, and different food items cause different worn shape to develop (browse maintains high and sharp features on the tooth surface, whereas grass “grinds” them down leading to them to progressively wear down lower and more blunted the more there is grass in the diet).”

hmnh-mammoth-and-mastodon-teeth

Examples of a mammoth tooth — used to eat mostly grasses and sedges — and a mastodon tooth — used to eat trees and shrubs. Notice the very different shape of these teeth for very different types of vegetation. Proboscideans such as mammoths and mastodons were once grouped in with ungulates, but this has changed. Picture taken at the Harvard Museum of Natural History by the author of this blog.

 

Obtaining data about the pollen record (non-arboreal pollen percentages, or NAP %) meant researching published information and connecting that information with the related fossil sites.

The mathematical work behind all of this–determining mesowear, animal body size, and then relating this to the available pollen record—is staggering.

Surely, I thought, isotopic analysis would have been a much easier way to obtain information about each fossil’s diet at least.  Especially given that the pollen record isn’t always available, or—in one case—runs the risk of being skewed by the defecation of Pleistocene hippopotamuses that grazed in the area.  Why, I wondered, did they rely on methods that seemed considerably more labor-intensive and potentially (to my understanding) less accurate?

“There are a number of reasons for this,” Juha explained. “First, we wanted to obtain as much palaeodietary data as possible, comprising as complete ungulate communities as possible, and this meant dealing with very large samples of fossil molar teeth. Taking isotope samples from all those teeth would have been laborious, time consuming and expensive, not to mention also slightly destructive to the fossil specimens.

cervus-elaphus-richmond-park-london-uk

Cervus elaphus (Red Deer, Artiodactyl) at Richmond Park, London; photo courtesy of Juha Saarinen. Red Deer are one of the most extensively studied animals today. You can read about another study that references Red Deer in this post.

 

“Second, stable isotopes work best at resolving herbivore diet compositions in tropical areas where carbon isotope composition reflects roughly the proportions of C4/C3 –photosynthesizing plants (roughly grass vs. browse) in diet, but outside tropical areas all plants, grasses included, are C3 photosynthesizing and the carbon isotope composition varies also considerably according to so called canopy effect (open vs. closed environment), not just according to diet, and thus isotopes would not have allowed us to estimate the amount of grass vs. browse in the Pleistocene European ungulates as consistently and quantitatively as we could with mesowear analysis.

“Third, mesowear has been specifically shown to reflect average grass vs. browse compositions in the diets of ungulate populations, without being significantly obscured by other environmental variables, such as climate or environmental openness (e.g. Louys et al. 2012, Kaiser et al. 2013). Even if mesowear is a ‘physical observation’ as you say, it has been shown to specifically reflect the amount of abrasive dietary items (mostly grass) in herbivore diets.”

The authors focused on fossil-rich sites, where they could study between 3 – 10 fossils of each species.  They made sure to include species that were browsers, grazers and mixed-feeders.

figure-1-saarinen-et-al

Screenshot of Figure 1 from “Patterns of diet and body mass of large ungulates from the Pleistocene of Western Europe, and their relation to vegetation.” Palaeontologia Electronica19.3.32A: 1-58

 

“I owe thanks to my co-authors who knew much of the available European Pleistocene mammal collections already, having experience on working on them for many years,” Juha responded when asked how they knew of or had access to so many fossils.

Adrian Lister from the Natural History Museum of London in particular has a huge amount of knowledge and experience about Pleistocene mammal collections.

“I was also in contact with the curators of the museum collections, who gave me valuable information about the how much and what kind of material they have. Also, information about important fossil finds and numbers of specimens found have often been published before in scientific journals.

“The authors of this paper represent different fields of research experience on the various aspects of the study. I started to work on this research as a part of my PhD work, and I originally planned it with my PhD thesis supervisors Mikael Fortelius, Jussi Eronen and Heikki Seppä from the University of Helsinki.

“During the work, I visited the Natural History Museum of London, where I worked together with Professor Adrian Lister, whose expertise on British Pleistocene mammals, the NHM fossil mammal collections and mammal palaeoecology in general were very important for this work.”

NHM-DrListerLyuba

Image of Professor Adrian Lister, Natural History Museum of London, with the mummified baby mammoth, Lyuba; photo courtesy of the Natural History Museum of London for this post.

 

This work was not without its challenges.  As with any study of fossils, there are limits to the number of fossils available.  While pollen record availability has increased, there is still so much more to be discovered.  And although some species–based on extant examples–do not exhibit sexual dimorphism in body size, the sex of most of the fossils they studied was indeterminate.

“Indeed, these were some of biggest challenges in this study,” Juha acknowledged, “but they were expected and nothing much could be done to completely avoid them. I would add that it was often challenging to connect the fossil mammals with associated pollen records, especially when the fossil pollen was not obtained directly from the mammal fossils. To succeed in this study, it was important to analyze lots of data in order to overcome these problems, and to ensure that the main results and conclusions of this study are robust despite of them.”

The authors of this paper considered numerous variables in their research, and they suggest that ungulate size has a lot to do with a number of factors.  This might seem obvious, but such has not been the result of past studies.  In particular, Bergmann’s rule, which stipulates that body size corresponds largely to thermoregulation (i.e.: big body size is the result of living in colder environments), has been supported before.

bison-bonasus-kraansvlak-netherlands

Bison bonasus (Artiodactyla), Kraansvlak, Netherlands;photo courtesy of Juha Saarinen. 

 

“[T]here has been a lot of discussion as to what ultimately explains the tendency of some (but not all) organisms to be larger in cold climate. This was actually one of the main questions I discussed in my PhD thesis,” wrote Juha. “Already in 1950s some researchers (e.g. Scholander 1955, Irving 1957, Hayward 1965) pointed out that increase in size alone would not give a large enough benefit for thermoregulation in cold climates, especially considering that mammals have far more effective mechanisms of keeping warm, such as thick fur.

“Since then, many authors have noted that while there is a tendency of mammals being larger in higher latitudes, there are a number of exceptions to this ‘rule’ and heat conservation alone would not explain it.

“However, body size in mammals does correlate with food quality and availability and this seems to explain most of the body size patterns observed in mammals (e.g. Rosenzweig 1968, Geist 1987, Meiri et al. 2007, McNab 2010). For example, many herbivorous mammals tend to be larger at higher latitudes because food quality is better there (e.g. because of fertile soils created by glacial erosion and because plant defense mechanisms are lower), and thus predators eating them also tend to be larger there, but for example brown bear body mass does not correlate with latitude but with distance to nearest salmon spawning areas. On the other hand, population density also affects body size through resource availability: individual body size has been noted to decrease in many species of mammals when population densities are high leading to increased intraspecific resource competition (e.g. Wolverton et al. 2009).”

The authors of this paper argue that environment–climate, open or closed vegetation, food availability and quality–and species social structure–large or small herds–affect body size.

“[T]here are many (often interconnected) factors which together affect body size,” Juha explained. “This makes it quite complicated and challenging to study what ultimately regulates body size in mammals (and other organisms).

“In fact, our results do not support Bergmann’s rule as such, because even if our analyses show that larger sizes seem to occur in some species in open environments, this is not because of low temperature, as some of the open environments were in fact quite warm. Also, we often see that when one species was particularly large in an environment, another species was particularly small under those same conditions. E.g., we found out that red deer (Cervus elaphus) tends to be large in open environments, but wild horse (Equus ferus) tends to be small in those same environments. Thus, our results do not support the assumption of Bergmann’s rule or any other “single-cause” explanation for ungulate body size variation.

“What ultimately regulates ungulate body size is primarily food quality and availability, which is affected by the interplay of vegetation structure (regulated by environmental temperature, precipitation and soil fertility), interspecific resource competition (depending on the presence of competing species) and intraspecific resource competition (depending on population density). For example, species with large population densities in open environments, such as reindeer, bison and wild horses, could be small under those conditions because of increased intraspecific resource competition, whereas species with smaller population densities in open environments, such as red deer are large under such conditions, e.g. because of abundant, high-quality food and diminished plant defense mechanics. This is also the main conclusion concerning our results of Pleistocene European ungulate body size variation.”

“I think that studying how mammals in the past interacted with their environments is important for understanding how these interactions work in general,” he concluded. “At present, environments and their mammal faunas are so heavily influenced by human activities, and they have lost so much of their original diversity, that I believe that we simply need to study fossil mammals and their palaeoenvironments to better understand how these things have worked and ‘should usually work’ in nature.”

equus-ferus-mongolian-wild-horse-lippeaue-germany

Equus ferus (Mongolian wild horse and Perissodactyl), Lippeaue, Germany;photo courtesy of Juha Saarinen. 

It was a great honor and pleasure connecting with Dr. Juha Saarinen!  Reading this paper and gaining more insight about it from him was absolutely fascinating!  An enormous thank you to him for all of his generous help!!

Additionally, Dr. Saarinen was extraordinarily kind and helpful in clarifying points about the research that I had misunderstood.  That is always appreciated.  THANK YOU!!

Reference:

  1. Saarinen, Juha, Eronen, Jussi, Fortelius, Mikael, Seppä, Heikki, and Lister, Adrian M. 2016. Patterns of diet and body mass of large ungulates from the Pleistocene of Western Europe, and their relation to vegetation. Palaeontologia Electronica 19.3.32A: 1-58 palaeo-electronica.org/content/2016/1567-pleistocene-mammal-ecometrics
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Persistence Cave: A rich resource for paleontological research

Caves whisper exploration and discovery.

Anyone who has ever set foot in a cave of any size cannot help but wonder what lies beyond, what lurks in the crevices, the darkness.  Stepping into a cave is stepping into the entrance of mystery just waiting to be revealed.  In a world that has been largely tamed to fit the human species, there are few spaces that still hold an element of danger.  These unknown spaces beckon to the adventurous: “Explore me!” And who wouldn’t answer that call?

Me, that’s who. I am perfectly happy learning about the discoveries in caves from other people, thank you very much.

For people like me, Twitter and blogs have provided tantalizing glimpses of such explorations the world over.  And one of the more fascinating adventures has taken place at Persistence Cave, just one cave of many at Wind Cave National Park, South Dakota.

“Wind Cave National Park is full of fossils. Almost everywhere you go there’s going to be fossils: in the cave and at the surface. So Wind Cave National Park actually has [perhaps] 30-40 fossil sites.”

PhD student Jeff Martin explained more about the work he and his colleagues conducted there last season as he and his wife were literally driving to Texas to begin a new chapter in their lives. He was in the moving truck; his wife was in the jeep ahead.  Jeff and I had been in touch by email from time-to-time over the past year. As luck would have it, and thanks to his seemingly unending generosity, the time to discuss Persistence Cave by phone was while he was on the open road.

Wind Cave—as we know it now—was named because of the air that blows through an opening within.  It was considered a sacred place to the Native Americans long before settlers knew of its existence.  The Lakota people refer to the Black Hills (where Wind Cave is located) as ‘He Sapa’, (although it is listed as ‘Paha Sapa‘ on the Wind Cave National Park site).  Eventually, in 1903, it became the 8th National Park, but the first one to center around a cave.

Persistence Cave, a much smaller and less-explored cave in the park, was discovered by accident by Marc Ohms, spelunker and physical science technician for the park, in 2004.  His initial foray into the cave was brief: moving a cap rock, peering inside, seeing a rattlesnake, and deftly removing himself from the opening.

But its value as a fossil site was discovered thanks to another member of the park.

“Rod Horrocks, Wind Cave National Park Physical Scientist, in 2013, collected some sediment for preliminary analysis to see whether the site is paleontologically productive,” Jeff explained by email earlier.

It was, and this analysis is what eventually brought several scientists from diverse locations together.

Rod Horrocks sent the material to Dr. Jim Mead, Persistence Cave Project Leader, then at East Tennessee State University, where Jeff was a Master’s student at the time.  Jeff eventually moved to the University of Maine for his PhD, where Dr. Jacquelyn Gill was his advisor.

Sharon Holte, PhD Candidate at the University of Florida, was also a previous Master’s student of Jim’s, as well as Dr. Chris Jass at the Royal Alberta Museum,” wrote Jeff, explaining the connections between the Persistence Cave teammates. “He knows that we each excel in different aspects of vertebrate paleontology, and he invited each of us to collaborate on [and] bring our expertise into the research project. I brought Dr. Gill with me to the Black Hills to see the cave and to learn how a paleontological excavation is usually conducted. She brings a different set of skills related to paleoecology and palynology.”

Also on the team are undergraduate Chason Frost from the University of Maine who studies horticulture.  His skills and those of Dr. Gill help the group understand that fossil plants and pollen found in the cave.

Sharon Holte, aside from being one of the three principal spelunkers in this dig, is in charge of educational components.  Chris Bell at the University of Texas Austin studies the fossil rodents; Dr. Chris Jass and Dr. Jim Mead study fossil rodents as well, but include fossil snakes.

“Each person has their role,” he said, “their own ecological-niche, if you will.”

And Jeff?  He is the “bison guy.”

“My PhD research and dissertation focuses on bison body size adaptation to climate change over the past 40,000 years and how does that evolutionary legacy influence the bison we ranch today,” he wrote before he graduated this past Spring. “To answer this, I am using Persistence Cave and other fossil sites in Wind Cave National Park boundaries to geographically isolate my variation to only local animals.”

Wind Cave National Park, currently home to 400+ extant bison, offers information on both fossil bison and their living descendants.

 

EPSON DSC picture
EPSON DSC picture; bison at Wind Cave National Park, public domain from the National Park Service

 

“Collectively, we (Jacquelyn, Chason, and I) will then also look at the pollen grains and macro-botanicals preserved in the sediment to reconstruct the paleoecology and paleoclimate of the Black Hills through the last 11,000+ years to today. This is [to understand] the climate and ecology the bison were living in at these times.”

But let’s get back to the cave itself.

Below is an image of Natural Trap Cave (another exciting fossil cave dig in Wyoming; photo from myfossil.org):

 

Natural Trap Cave from myfossil.org

 

Compare that to an image of Persistence Cave from the top looking in (photo: Chason Frost as posted on Jeff Martin’s blog here):

 

Photo by Chason Frost - Persistence Cave entrance from Jeff's blog

 

 

 

And one of Sharon Holte peering out:

 

CB - SHolte peering out of cave

 

 

Finally, below is an image from the Rapid City Journal of “a tight spot in Wind Cave” (photo: National Park Service):

Marc Ohms WCNP National Park Service

 

When I asked about how this image compares to the space within Persistence Cave, I was surprised by Jeff’s email response.

“The picture above is much larger than the cave we are working in,” he described of the 2015 dig.  “The cave is very narrow and only fits one person’s shoulder width and up to 1.5 shoulder widths in places. The vertical height is similar to the above photo though.”

“I’m a broad shouldered fella’ and very, very tall,” he continued by phone recently. “The space in there to turn around is not quite enough for me, so I’d have to climb in and then climb backwards out.”

“Chris Jass and I are both the exact same height. Chris is a far more experienced spelunker, and even Chris wasn’t going in there.”

Sharon Holte, Chason Frost and Jim Mead were the principal spelunkers for the site.  Only one person could be in the cave at a time, and their only source of light came from a headlamp.  Trowels, buckets and ropes: their only tools.

 

CB - Sharon Holte important gear


“I thanked them endlessly, and I still thank them for all the work they were doing down in there,” Jeff said of his three colleagues. (A video of Sharon’s work in the cave can be found here.)

Work involved taking chunks of sediment in buckets out of the cave, tagging it, labeling the information (where that sediment appeared on the appropriate grid, at what depth, etc.), bagging that sediment, and then sending it down—by zipline, of all things!—to the truck below, where it could be taken to be screenwashed by other team members. (You can see a video of that process here, on Jeff’s blog.)

 

CB - screenwashing for microfossils

Screenshot of tweet during the 2015 Persistence Cave (#cavebison) dig

 

Their fossil discoveries have been diverse. Jeff wrote that “[a] camelid, (the species is unknown at this time), has been an extraordinary find. We have 5 different kinds of snakes and at least 5 different species of bats. [A] pika is also an intriguing find.”

 

 

CB - Jim Mead and snakes

CB - fossils found

 

CB - snake fossil

 

CB - toe bone and Jeff Martin

 

CB - Jeff Martins favorite bone found at that point

Screenshots of some of the many tweets during the 2015 Persistence Cave (#cavebison) dig

 

“One of the fun things that we ran across was a ton of Ponderosa pine needles,” he mused later by phone. “That’s the primary tree out there now.  Today, they’re mostly a two-needle bundle. In the past, it seems as though they were a three-needle bundle. And we don’t know exactly what that means yet.  So we’re trying to figure out if that means anything at all; if it’s a genetic difference; or if it truly is an environmental difference that it’s responding to.”

 

CB - Twitter conversation about plants

Screenshots of some of the many tweets during the 2015 Persistence Cave (#cavebison) dig; the scientists involved in this dig didn’t just conduct research, they also conducted outreach to the larger public through social media.

 

 

Work did not continue as expected on the site this year for a number of reasons, but it’s not over yet.  Studies on the fossils continue at the University of Maine (pollen and plants); the bison fossils have travelled with Jeff to Texas A&M University where he is now in wildlife sciences; and the rest of the fossils are housed at The Mammoth Site, where Dr. Jim Mead is currently Chief Scientist and Director.

The Mammoth Site is another major connection between many of the team members, as they have each “worked [there] at some point…over the last 40 years.”

As many know, that site is a paleontological (and proboscidean!) goldmine turned museum, thanks to the work of many, including the late Dr. Larry Agenbroad.  Over 60 mammoth fossils have been discovered there to-date, among other fossil species.

Bonebed at The Mammoth Site

Image of the bonebed at The Mammoth Site where excavations continue to this day

 

“He was probably THE reason that I got into the School of Mines [as an undergrad] and was also the reason I got into paleontology,” Jeff said of Dr. Agenbroad.

“I’m not alone,” he continued. “There are several of us that are like that.  We all stem from Larry.”

The reverence in his voice was not difficult for me to understand.

Jeff’s introduction to this paleontologist began when he was much younger, through the 2000 documentary “Raising the Mammoth.” The film focuses on the Jarkov mammoth, and Bernard Buigues’ attempts to excavate it.  The team Buigues calls upon to help include some giants of proboscidean research: Dick Mol and Larry Agenbroad.

A year or so after seeing that film, Jeff’s family traveled to The Mammoth Site.  It was winter in South Dakota, and, he said, his family basically had “the run of the whole place.”  With a graciousness I am sure permeates everyone who works at that site, one of the interpreters (‘docents’) offered to bring Dr. Agenbroad out to meet them.

“There’s 8-year-old me that’s just giddy with joy to be able to meet one of my idols,” Jeff shared with no small amount of enthusiasm. “And then he said, ‘You’re a little bit too young to work for me. Come back when you’re older.’”

“So that’s exactly what I did. I worked for him in [the summers of] 2007 at the Hudson-Meng Bison Kill Site and  2008 and 2009 at the Mammoth Site as an intern while I was at the School of Mines.”

Dr. Agenbroad passed away two years ago, followed by his wife, Wanda, a month later.  This saddened me as someone who did not know him closely; I could only imagine how this affected Jeff, who had.

“I’ve made my peace with it,” he acknowledged, and then said something that truly moved me: “I have several things that Jim [Mead] gave me…and one of them is a pocketknife that I carry on me every single day. One of the same pocketknives that Larry carried on him every single day. So I’ve got Larry with me, right now, as a matter of fact.”

Jeff and his colleagues hope to resume work at Persistence Cave next year.

As we discussed some of the findings from last year’s dig, he said, “The oldest date right now at Persistence Cave is at 39,000 and the youngest date is at 3,200.  We have some 37,000 years of deposits with bison throughout. And we also have [modern-day] bison living at the surface!”

Jeff’s research, both of Persistence Cave and of Project Bison, underscore his passion for this animal, as well as the desire to understand its ecological significance.

“I’m looking at both the fossil record and looking at their body size, using the calcaneum [heel bone] as the proxy for body mass. And then also comparing that to modern bison that have just recently passed away within the past 1-3 years.  That’s what I was doing this past summer: going to carcass sites and measuring their calcanea. The unique thing about Wind Cave is that they have almost every single animal microchipped. So they can track this animal throughout its life. On top of that, they bring them in once a year and weigh them. So now we have a known mass of these animals and now a known measurement, because I measured some of their calcanea.

“I’ve got some [fossil bison calcaneal] measurements that go up to 180 millimeters, and I also have Bison bison today that the longest that I’ll find are 130 millimeters.  So quite a body size change in between the fossil and modern.”

Jeff presented some of his research at last year’s Society of Vertebrate Paleontology (SVP) meeting in Dallas.

Describing the results, he explained, “As it gets colder, bison get bigger.  As temperatures are increasing, bison get smaller. That has modern day application to the bison industry today. If we’ll have smaller bison with future global warming, we’re going to have to change our management options.”

As I pondered all of the information Jeff had shared with me about the work he and his colleagues had done, I couldn’t help but go back to the images of how small the cave actually is. If Wind Cave National Park has an abundance of fossil sites, why go through the trouble of trying to access this one?

“Surface localities often represent a one-time event,” he explained. “Persistence Cave represents many events over a long period of time. That’s the unique part of this locality.”

I will continue to enjoy their adventures from the safety of my computer!

 

**************

Jeff Martin: you were extraordinarily generous with your time and responses to my myriad questions.  Likewise, I am in awe of how open you were with your experiences.  For being willing to share all of this, I am truly grateful.  It was an honor and a pleasure connecting with you!

When #CaveBison starts up again, you can be sure it will be on Twitter!  Follow these scientists:

@BisonJeff

@JacquelynGill

@SharonHolte

@Pocket_Botanist

@MammothSite

 

You can follow Jeff’s research here and here

Jacquelyn Gill is one of three hosts of the podcast, Warm Regards, which discusses climate change.

 

Meet Dr. Katy Smith – Mastodon Detective

If you imagine the Great Lakes region over 10,000 years ago, you might see large, hairy beasts with relatively straight tusks grazing around boggy areas or moving within dense forests.  Their fur and overall appearance might cause you to confuse them with woolly mammoths, but these are the mammoths’ shorter, stockier cousins.  And if any of them would let you get close enough to inspect their mouths, you’d see in an instant that their teeth are completely different than those of mammoths.

 

[image of contemporary boggy area in Alaska, courtesy Getty Images]

 

Whereas mammoths are believed to have eaten grasses and even flowers, mastodons needed teeth suited to the mastication of hardier stuff: shrubs, parts of trees, perhaps pinecones?   Mastodon teeth, with the bumps and ridges one might associate with carnivores, are easily recognizable as ‘teeth.’  Mammoths, in contrast, needed to grind food, producing teeth with spherical lengths of ridges across each tooth.

ISM - Mastodon tooth

 

[image courtesy of Ron Richards, Indiana State Museum, for this post: Mammoths and Mastodons in Indiana – Part 1.  Can you tell which tooth belongs to which species?]

 

ISM - Mammoth tooth

 

[image courtesy of Ron Richards, Indiana State Museum, for this post: Mammoths and Mastodons in Indiana – Part 1.]

And while woolly mammoths pervade popular culture and interest, there are some, like Dr. Katy Smith, Associate Professor of Geology at Georgia Southern University and Curator of the Georgia Southern Museum, who prefer their lesser-known cousins and have made fascinating contributions to our understanding of them.

Mastodon discoveries usually produce the fossils of a single animal, and rarely a complete skeleton. Rarer still, finding skeletal remains of multiple mastodons at the same site.

Such a unique discovery occurred in 2005, when more than 300 fossils were found in Hebron, Indiana.  Now known as the “Bothwell site,” it was originally going to be the location of the landowner’s pond.  Instead, Indiana State Museum paleobiologist Ron Richards and his crew uncovered bones that included numerous mastodons (Mammut americanum), giant beaver (Castoroides) and hoofed animals with even-toes (artiodactyls).

ISM - 2005 Bothwell Mastodon 2

 

ISM - 2005 Bothwell Mastodon 1

[images of the Bothwell site dig, courtesy of Ron Richards, Indiana State Museum, for this post: Mammoths and Mastodons in Indiana – Part 2.]

 

Four years later, the Bothwell site became the focus of Katy Smith, her dissertation, and two subsequent papers she co-wrote with Dr. Daniel Fisher at the University of Michigan.

But let’s take a moment to consider what paleontologists uncover. However rudimentary this may seem, it is important to note that bones are generally not discovered in neat order, intact and with each skeletal component attached where it would have been in the life of the animal.

Consider, too, that not all bones survive.  And those that do are often broken or in terrible condition.

So even at a site such as Bothwell, which produced lots of fossils, a paleontologist’s job is no less challenging.  The pieces of information are incomplete, mere clues to the animals that died there.

The questions, however, are profuse.

Why were so many animals found in that one spot?

If, as it is currently debated, mastodons shared behavioral traits with modern-day elephants, was this a family unit?

If so, was this group—like elephants–comprised largely of female and juvenile mastodons?

And why were other unrelated animals discovered among them?

Did a sudden disaster kill them all?  Were humans involved?

 

Embed from Getty Images

 

Sexual dimorphism is another way of referring to the traits that make an animal either female or male.  Some of us would assume, since mastodon pelvic bones were not among the Bothwell fossil assemblage, that the sex of these animals would remain unknown.

There were 13 mastodon tusks, only four of which were complete. And this, remarkably, is what prompted Katy Smith’s research.

“I wanted to know if I just had tusks, what can I do to figure out if I’m looking at a male or a female,” she explained by phone.

Katy Smith - measuring an African elephant tusk

 

[image of Dr. Katy Smith measuring an African elephant tusk in (what this author believes must be one of the greatest places on earth) the basement and fossil collection of the University of Michigan; courtesy of Dr. Katy Smith]

 

“Other people have looked at [sexual dimorphism], but I wanted to look at it specifically with the Bothwell mastodons, because they were inferred to be female, and female mastodons are less common in the fossil record than males.

“When I presented preliminary results from my research in a paleontology class, the professor said, ‘Why don’t you try multivariate analysis?’ And it just kind of spiraled from there.”

Multivariate analysis,’ as the name implies, means using more than one type of measurement or observation towards a hypothesis.  In other words, rather than simply using size as a determination of sexual dimorphism, applying numerous methods and statistics that support or disprove it.

Already, the amount of information scientists have pulled from tusks alone is fascinating.

Tusks are teeth.  They are described, in Dr. Smith’s dissertation as “hypertrophic incisors.” And, unlike human teeth, they continue to grow the entire life of the animal. So where we can simply look at a human tooth and know immediately whether it is from an adult or a child, the same cannot be done with tusks.

What their hardy structure records includes the age of the animal, growth in winter or summer months each year, their overall diet, and periods of nutritional stress.  (As described in an earlier post, Proboscidean molars can even provide details regarding where they roamed during life.)

But much of this information can only be gleaned from well-preserved, intact tusks, as well as from cutting into and examining their chemical composition.

“If you don’t know what the sex of the animal is before you look at tusk microstructure,” she said, “it can be hard to interpret what you’re looking at.”

Part of what Dr. Smith hoped to discover were similarities in the tusks where sex and age had already been determined.  If certain structural elements were the same across female mastodon tusks, such that they tended to differ from male mastodon tusks, this might help determine sexual dimorphism in future tusk discoveries.

She also hoped to discover any similarities between the tusks of extant elephants and mastodons.

Katy Smith -longitudinally bisected tusk

 

[image of longitudinally bisected tusk, courtesy of Dr. Katy Smith] 

 

Thus, she studied and measured tusks of both species from numerous museum collections. (Asian elephant tusks were not used, as female elephants of this species tend to have either tiny tusks or no tusks at all.)  She rather amusingly refers to the approximate amount of tusks involved as “5,000 pounds of tusk.”

Her dissertation and the two papers describe the type of analysis performed in detail.  Among them were canonical variates analysis (CVA) and discriminant function analysis (DFA).

“Fortunately, we didn’t have to cut into the tusks to do those measurements. You just insert a stiff wire into the pulp cavity.”

“We think about tusks sometimes as stacks of sugar cones, because they actually grow in a kind of [layered] cone structure. So you think about one sugar cone, and then you put another one inside that one and then another one inside that one and so on and so forth. And the last sugar cone is empty. There’s nothing in it. That represents the pulp cavity.”

“[Analyzing the] pulp cavity is probably one of the best single measurements that you can use to distinguish between male and females. [I]n females, that pulp cavity will terminate before the gum line, and in males, it will terminate after the gum line, closer to the tip.

“This is something that we saw in almost every mastodon. So it was kind of cool.”

 

Katy Smith - female mastodon

 

[image of female mastodon skull and tusks, courtesy of Dr. Katy Smith]

 

“If we could have cut every tusk, I would have,” she admitted, and laughed. “But it was a matter of collecting these measurements at different museums. And so I would just go there and collect all of them, and that was how we’d get the pulp cavity depth.”

“I’ve always been interested in paleontology,” she said when I asked her how she got started.

“I’m one of those kids who just never grew out of it. My parents used to take me to the museum all the time, and I used to spend hours and hours staring at the dinosaur dioramas there, just loving it.  I told my kindergarten teacher I wanted to be a paleontologist. I never changed! My 5-year-old self grew up and became a paleontologist.”

But her interests moved away from dinosaurs when she realized that their fossil record in Wisconsin, her home state, was rare to nonexistent.

After all, she said, “I started just wanting to explore what was underneath my feet.”

It wasn’t until grad school at Michigan State, where she met the late Dr. Alan Holman, that she realized her passion for mastodons.  His own interest in the species was infectious, and it was through him that she learned of the numerous mastodon (Mammut americanum) fossil discoveries in the area.

“Wow!” she said, recalling her initial reaction. “There are over 300 mastodons in Michigan. This is exciting!”

Katy Smith - male mastodon

[image of male mastodon skull and tusks, courtesy of Dr. Katy Smith]

Not surprisingly, she did her PhD work at the University of Michigan, home to Proboscidean expert Dr. Daniel Fisher, who was her advisor.

“I wanted to work with him,” she explained, “because I wanted to continue working on mastodons, and he had a couple of ideas for projects. One of them included this assemblage of mastodons from Indiana, which were—supposedly—all female.”

What she discovered regarding the Bothwell site is both thought-provoking and fascinating:

  • 8 tusks were determined to be female; the other 5 are unknown
  • the ages of the mastodons range between 19 and 31 years old
  • there is evidence that at least one juvenile might have been among them (a “juvenile tooth crown” was found)
  • given that two mastodons died in winter, and another two died either in late summer or early autumn, this indicates that the collective deaths of these animals didn’t happen at the same time (hence, not a single event)
  • none of the mastodons appeared to be under nutritional stress when they died
  • members of a family unit would be expected to have the same “isotope profiles”–chemical signatures in their teeth–but these do not

Based on the evidence provided, Dr. Smith wonders whether these animals were part of a meat cache for humans (members of the Clovis culture) that co-existed at that time.

But perhaps the single most remarkable result of her research is helping other paleontologists–who often have nothing more than a single tusk–determine the sex of that animal using her different types of analysis.

Prior to her dissertation, only one female mastodon tusk had been analyzed for growth rate.  To date, I am unaware of any other publication (paper or book) that helps detail the sexual dimorphism in mastodons by tusks alone.

When I remarked upon this, I asked her if others had cited her work.  Her response, after stating that others had, was equally fascinating to me.

“It’s always the hope as a scientist that you’re contributing in some way,” she said, “and you know that you’re contributing if somebody else is using what you’ve done.”

 

An enormous and sincere THANK YOU to Dr. Katy Smith for her generous and fascinating answers to my many questions, her gracious help when I had trouble understanding certain points, and for being so much fun with whom to connect! I cannot express how much I wish I could attend her classes, nor how fascinating I found her dissertation. I am profoundly grateful that she shared it with me!

A sincere thank you to my Dad, as well, for helping me understand tooth components (i.e.: dentin, cementum)!

**A quick reminder that I am neither a scientist nor a paleontologist, so any errors in this post are my own.

Bothwell Mastodont Dig, courtesy of Indiana State Museum; many thanks to Bruce Williams and Leslie Lorance!

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References:

 

Other references:

 

Cohoes mastodon size comparison

[image of sign in the NY State Museum illustrating the size difference between an extant elephant, a woolly mammoth and the Cohoes mastodon; picture taken by the author]

Exciting New Info About Mastodons and Humans – Yukon Paleontology, Part 1

“Good morning!”

It’s not just a greeting; it sounds like a proclamation.

The voice on the other end of the phone is deep, melodic, and—as our conversation progresses—punctuated with moments of laughter.  We have been discussing paleontology in the Yukon, and with each new detail, I begin to wonder why this territory is not making regular international headlines.

Dr. Grant Zazula’s work is fascinating, and it is neither a short phone call nor the only communication we’ve exchanged. And yet, it is all that I can do not to encourage him to keep going, long after social decorum dictates that he has been more than generous with his time.

Dr. Zazula and mastodon leg

[image of Dr. Grant Zazula with a mastodon ulna, part of the Earl Bennett mastodon, courtesy of the Government of Yukon]

Dr. Zazula is the Yukon paleontologist, a job that has only existed since 1996. His own tenure began in 2006.  With an office in Whitehorse, the capital of the territory, his work oversees an expanse of Canada that abuts Alaska.  It is a land of dramatic beauty, where colors dance in the sky and mountains tower in silent grandeur.

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His most recent paper, co-written with 14 other people, made news throughout the world and continues to attract media attention. In it, the scientists present data that completely overturns previously believed information about extinct animals and the impact that humans may or may not have had upon their survival.

“[T]here were two radiocarbon dates in the literature from Yukon mastodons,” he explained in an email. “One that was ~18,000 and the other 24,000 years old.”

“Based on analysis of the paleoecology, that was a time when steppe-tundra grasslands covered Alaska, Yukon and Beringia. There were probably no trees, few shrubs and almost no standing water. It was very cold and, especially, dry. This seemingly is not good mastodon habitat. So either the dates were incorrect, or our understanding of mastodon ecology, behavior and adaptations need[s] to be revised.”

Various species of mastodon once existed throughout the world.  Although their fossils look elephantine, they are not believed to be direct ancestors of today’s elephants. They are, however, part of the same umbrella mammalian group: the Proboscidea (so-named for the trunks possessed by many—but not all–of their members).  In North America, that group contained the American mastodon (Mammut americanum), the woolly mammoth (Mammuthus primigenius), and the Columbian mammoth (Mammuthus columbi).

Cohoes mastodon

 [image of the Cohoes mastodon, NY State Museum, Albany; taken by the author]

Mastodons tended to have straighter tusks and were shorter than their mammoth cousins. They also ate hardier vegetation, food that required a much different tooth structure than mammoths.

ISM - Mastodon tooth

[image of mastodon tooth, courtesy of the Indiana State Museum]

ISM - Mammoth tooth

[image of mammoth tooth, courtesy of the Indiana State Museum; for more info about the differences between mammoths and mastodons, see this post.]

Parts of Siberia, Alaska and the Yukon were once connected in an area known as “Beringia.”  The Bering Strait did not yet exist, enabling animals and eventually the first humans to cross into our continent.  It is believed that humans arrived in what is now North America about 14,000 years ago.

And this is where the research of Dr. Zazula and his colleagues becomes particularly important.

Prior to their paper, one theory to mastodon extinction laid the blame upon first humans: it was proposed that they overhunted these animals.

Sampling 36 fossils and presenting 53 new radiocarbon dates, Dr. Zazula and his colleagues found that mastodons within Alaska and the Yukon were much, much older than the originally published dates.  In other words, their research suggests that mastodons from what was once Eastern Beringia were no longer present when the first humans appeared.

The path to this remarkable research did not happen overnight.

The foundation appears to have been laid by two different events: by the chance meeting of Dr. Zazula and a gold miner, and later, by the PhD work of a graduate student.

If one reads the acknowledgements on the aforementioned paper, Dr. Zazula references Earl Bennett as both the donor of a partial mastodon skeleton and his inspiration to learn more about mastodons within the Yukon.

“Earl is a great Yukoner,” Dr. Zazula wrote when asked about this. “He mined for gold underground in the winters with a pick and shovel, decades ago. He worked on big gold dredge machines. And, he loves paleontology.

“While mining, he made collections of Ice Age bones that were just left around the mining camp or were encountered while mining. He eventually amassed an amazing collection.

“In the early 1970’s a gold dredge on Bonanza Creek hit a skeleton of a mastodon. An incredibly rare find! Someone collected it and was looking to sell it. So, Earl bought the skeleton just to make sure that it never left the Yukon. He had it in his garage for decades.

“One day a mutual friend introduced me to him in a coffee shop, about a year after starting my job [as the Yukon paleontologist]. He said that he had a mastodon skeleton and wanted me to see it. I ‘corrected’ him, saying that it was more likely a mammoth, because we almost never find mastodons in the Yukon. He assured me he know the difference and said he would see me tomorrow at my office.

“The next day he backed his truck up and in it was a partial mastodon skeleton. I couldn’t believe it. There were several postcranial bones, some vertebra, scapula, parts of the skull and parts of the mandible with teeth. It was amazing. I wanted to find out how old it was, and that was one of the inspirations for this project. Earl is a good friend now and big supporter of our research.”

Bennett mastodon skeleton

[Paleontologist Grant Zazula with a partial American mastodon (Mammut Americanum) skeleton found on Bonanza Creek and donated to the Yukon fossil collection by Earl Bennett, from Ice Age Klondike, courtesy of the Government of Yukon]

That partial skeleton was indeed one of the many fossils sampled for the paper.

Dr. Jessica Metcalfe, one of the co-authors, also prompted this research when conducting work for her PhD.

“[S]he was doing a project looking at stable isotope ecology of mammoths and mastodons in various places in North America,” said Dr. Zazula.

Jessica Metcalfe with mammoth bone

[image of Dr. Jessica Metcalfe with mammoth bone, courtesy of the Government of Yukon]

Her work included Yukon fossils that were sent to the lab at the University of Arizona to be radiocarbon dated.   Those dates turned out to be older then 50,000 years old.

“So that’s what got me thinking,” he continued, “‘well, maybe those original published dates are wrong.’”

“The first step was to re-date [the specimens that had produced the original published dates]. The new dates turned out to be >50,000 years. So we knew there was a problem with the previous dates. We figured then we should date as many as we could get our hands on.”

This lead Dr. Zazula to connect with Dr. Ross McPhee, another co-author.

“I got in touch with him early because he oversees collections at the American Museum of Natural History, [and] he has a big interest and lots of experience working on Ice Age extinctions. [H]e’s an excellent writer and really kind of kept us going with some of the writings. He was really integral to keeping things together.”

The paper eventually involved a total of 15 people.

“I feel pretty strongly that if you worked on it and contributed to it, then you should be considered an author,” Dr. Zazula stated.  “So it ended up being a long list.”

One of the first aspects their paper addresses is the reason behind why the original published dates are incorrect: the dating analyses were contaminated by fossil conservation methods.

“Humic acids in soils can be absorbed by the bones and teeth and chemically bind themselves to the collagen,” he wrote, explaining further. “So, modern ‘young’ carbon in those acids basically contaminates the ‘old’ collagen in the ancient fossil. And, it can be tricky to remove.

“The same with consolidants in museums. Varnish, glue, and other substances to preserve fossils can be absorbed into the bone and chemically bind with the collagen in the bone. These substances probably contain young, modern carbon which messes up the radiocarbon dating measurements.”

When asked whether museums continue to use the same preservation products that contaminated the dates, he wrote, “Yes, for sure. The thing is now museums keep better records of what they use. Many of the fossils we dated were collected in the 1940’s or at least several decades ago. Museums were not that vigilant about keeping detailed records on those things then. Also, they seemed to put preservatives on everything. Now, at least if we know what was put on it, the chemistry can by developed to remove it. Most of the common preservatives now are soluble in alcohol or acetone and can be dealt with. The problem is when they are unknown.”

We discussed this further by phone.

“One thing about Alaska and the Yukon,” he said, “is that the Ice Age bones that come out of the ground are so well preserved because of the permafrost. In other localities, say, the deserts of the American Southwest or the Great Basin or the Plains, where bones have been out in the sun and [are] dry and hot, they [sometimes] fall apart really easily when they come out of the ground. They need to be glued and consolidated with these various types of museum products.

“So you kind of have to weigh the different values.

“Say if it’s a specimen that’s already been radiocarbon dated, and it starts to slowly disintegrate, well, then you kind of have to intervene or else you’re just going to end up with a box of dust and broken bone. You have to decide whether the importance is more with display or preservation of the morphology versus needing to radiocarbon date or other types of analysis.

“[Y]ou have to look at the pro’s and con’s of whether the sampling [for radiocarbon dating] will ruin the specimen or not, and what is the potential information that can be gained by doing it. To me, I feel that having a research collection [in the Yukon], it’s all about research and learning new things from these specimens.”

Ultimately, I wondered whether Dr. Zazula expected the results he and his colleagues uncovered.

“I wasn’t quite sure,” he answered. “I had the gut feeling that these previously published radiocarbon dates were probably wrong. It didn’t make a lot of sense ecologically to have mastodons living in the far North when it was seemingly habitat they couldn’t live in: habitat with grassland and cold, dry steppe tundra conditions, no trees and very few shrubs.

“But there [was] also a part in the back of my mind that thought, ‘well, if those [previously published dates] were right, that’s maybe even more interesting because they are telling us something about mastodons and their behavior and their adaptations that we didn’t know before.’”

————

It was a great honor and pleasure to connect with Dr. Grant Zazula! Not only patient with my myriad questions, he is an adept and fascinating ambassador for the Yukon. A Mammuthus columbi-sized thank you to him!

A Mammuthus columbi-sized thank you to Dick Mol, as well, who is the reason behind this post!

Dick Mol with horse skull

[image of Dick Mol with fossil horse skull, found near Dawson City, Yukon; courtesy of the Government of Yukon]

Yukon Paleontology Program: http://www.tc.gov.yk.ca/palaeontology.html

Articles and publication referenced:

 

Listen to Dr. Zazula discuss his paper on the CBC’s Quirks & Quarks: http://www.cbc.ca/radio/quirks/quirks-quarks-for-dec-6-2014-1.2864605/mastodons-made-an-early-exit-from-the-north-1.2864634