Researching Fossil Ungulate Communities

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.

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?

 

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).”

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 (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.

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.”

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 (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 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

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.

 

[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?]

 

 

[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).

 

[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.

 

[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.

 

[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.”

 

 

[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!”

[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!

—————

References:

• Life histories of female American mastodons (Mammut americanum): Evidence from tusk morphology, stable isotope records, and growth increments, Kathlyn Mai Smith, Dissertation, University of Michigan, 2010
• Sexual dimorphism of structures showing indeterminate growth: tusks of American mastodons (Mammut americanum), Kathlyn M. Smith and Daniel C. Fisher, Paleobiology, 2011
• Sexual Dimorphism and Inter-Generic Variation in Proboscidean Tusks: Multivariate Assessment of American Mastodons (Mammut americanum) and Extant African Elephants, Kathlyn M. Smith and Daniel C. Fisher, Journal of Mammalian Evolution, 2013

 

Other references:

• Woolly Mammoths’ Taste for Flowers May Have Been Their Undoing, NPR
• Fifty thousands years of Arctic vegetation and megafaunal diet, Nature, Eske Willerslev et al.
• Ancient Life of the Great Lakes Basin, J. Alan Holman, 1995, University of Michigan Press
• Mammoths & Mastodons of the Ice Age, Adrian Lister, 2014, Firefly Books Ltd.

 

[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]