Dr. Advait Jukar – Solving Mysteries in South Asian Fossil Communities

Dr. Advait Jukar–Deep Time – Peter Buck Fellow at the Smithsonian Institute–wants to really understand ancient ecosystems in South Asia, but doing so means beginning with some of the very basics. Challenges include not just a lack of available fossils from the region, but also the lack of detailed records from early paleontologists and a dearth of contemporary research.

He is, in a sense, an explorer.  (All paleontologists are.)  If we think of paleontology as a sculpture in progress, many scientists are working on the fine details.  Dr. Jukar, on the other hand, has the clay and the tools, but the sculpture itself hasn’t even begun to take form.

Consider what he has to work with: isolated proboscidean teeth and skulls, for example, collected by Hugh Falconer and his crew in the early 1800s.  They didn’t record where the bones were found, let alone where each fossil was in relation to the other.  Those who later described these fossils made dubious claims regarding the species.  And few people to this day have revisited this data or expanded upon it. 

 

Image of Dr. Advait Jukar at work with an Elephas hysudricus molar; courtesy of Dr. Jukar

 

Compare this to Maiasaura fossils in the Northwest US.  So many fossils of this species have been excavated that Dr. Holly Woodward Ballard has created the Maiasaura Life History Project.  Its goal is to uncover more details about this particular species than any other currently known extinct creature.  She has a wealth of data at her disposal. Unlike Dr. Jukar, the fossils she can study have been found fairly well articulated, very well documented, and in remarkable abundance.  There are adults, sub-adults, juveniles and embryos.  She and her colleagues are able to add to existing scientific literature using the latest technology.  It’s an exciting project with absolutely fascinating possibilities.

 

FIGURE 6. Survivorship curve for Maiasaura. Sample size of 50 tibiae was standardized to an initial cohort of 1000 individuals (assumes 0% neonate mortality). Survivorship is based on the number of individuals surviving to reach age x (the end of the growth hiatus marked by LAG x). Age at death for individuals over 1 year old was determined by the number of LAGs plus growth marks within the EFS, when present. Error bars represent 95% confidence interval. Mean annual mortality rates (μ^) given for age ranges 0–1 years, 2–8 years, and 9–15 years. Vertical gray bars visually separate the three mortality rate age ranges; courtesy Dr. Woodward Ballard for this post

 

 

 

But so, too, is Dr. Jukar’s intended research.   Focusing on the tail end of the Neogene, about 4 million years ago, through the Quaternary, he wants to understand herbivorous mammals—their community, their ecology, their biogeography.  It’s just a question of building the necessary foundation first.

“I started to compile all of these species lists,” he explained by phone, “and saw that there were lots of species of proboscideans in South Asia during that period of time.  We have gomphotheres; we have stegodons; we have elephants.”

One way to understand an animal’s impact on its environment is to assess its body mass.  How big (or small) were these animals?  And therefore, how much did they need to eat?  A larger animal would presumably need to eat a larger amount of vegetation.  Similarly, a larger animal might reproduce less frequently than smaller animals.  Body mass reveals clues about how an animal fits into the ecosystem.

However, he continued, “I hit a wall because there was no way for me to estimate how big these elephants were.  The problem was they were largely known either from skulls or teeth, and the traditional methods to estimate the weight of an extinct elephant were using shoulder height or the length and circumference of the long bones. So if I have a skull but I don’t have long bones, I’m sort of in a bind, because now I can’t estimate how much this animal weighed when it was alive.”

He looked to methods that others have used in the past. One method used by his colleagues at Howard University seemed to be a promising fit.  They used the occipital condyle breadth of seacows—a proboscidean relative–as an indicator for body mass.  Dr. Jukar’s PhD advisor, Mark Uhen, mentioned that this method had also been used on yet another large mammal: the whale. 

The occipital condyle is a bone found at the base of the back of the skull, connecting the skull to the spinal column.  It’s a relatively small bone.  Why would this have an impact on determining body mass?

“If occipital condyle breadth is correlated with the size of the animal,” Dr. Jukar said, “and if the occipital condyle is the point where the skull attaches to the rest of the skeleton, then maybe the size of the skull scales with the size of the overall body. And if that’s true, then maybe the occipital condyle breadth will scale with the size of the limb bones as well.”

 

Image of the back of a mastodon (nicknamed ‘Max’) skull at the Western Science Center in California displaying the occipital condyle bone resting on the metal stand; photo taken by Jeanne Timmons

 

In other words, if a paleontologist has but one skull of an extinct proboscidean and no other related fossils, can that person measure the breadth of the occipital condyle as a way to determine the size and weight of that animal?

To test this theory on proboscideans, he researched available scientific literature and visited a number of museum collections.  Ultimately, he and his two co-authors, S. Kathleen Lyons and Mark Uhen, compared the occipital condyle breadth to the length and circumference of leg bones within extant elephants and extinct relatives.  Two elephant species were studied, as were six gomphotheres, three mastodons and one stegodon.  

Image of a fossil Moeritherium at the Yale Peabody Museum; photo taken by Jeanne Timmons

 

While careful to note that this method has its limitations, the results were promising.  The equations are different for each proboscidean family (gomphothere body structure and size is not the same as that of a mastodon) and they do not work for some of the smaller proboscidean species, such as Moeritherium.  In layperson’s terms, this research works for taller, lumbering proboscideans, not those with much shorter limbs and a perhaps waddling gait. Their paper and its results were published in the Zoological Journal of the Linnean Society: A cranial correlate of body mass in proboscideans.

This, though, is just the tip of the iceberg in terms of Dr. Jukar’s research.  An enormous collection of fossils from India resides in the Natural History Museum of London.  Found in an area referred to as the Siwalik Hills (or the “Siwaliks”) at the base of the Himalayas, Scottish paleontologist Hugh Falconer and his team collected them in the 1800s.  Among them are several stegodon teeth and skulls. 

Image of Dr. Advait Jukar measuring a Stegodon ganesa fossil in the Natural History Museum of London collection; courtesy of Dr. Jukar

 

The two species of stegodon excavated from the Siwaliks are, to this date, known as Stegodon insignis and Stegodon ganesa.  The species have very similar teeth, but their skulls seem to differ greatly.  The skull of S. insignis, according to Dr. Jukar, is “almost triangular in shape with relatively small tusks,” which Falconer chalked up to sexual dimorphism.

“Which I just thought was the weirdest thing to ever say about stegodons because the skulls are clearly different. They’re clearly not sexually dimorphic.”

Moreover, there seems to be confusion regarding which fossils Falconer assigned to which stegodon species that continues to this day.  

“So what was going on in his mind? I have no idea.  It’s a problem! Because since then, people have said that both of these must be the same species without really truly investigating them. 

“Any Stegodon tooth that they’re finding in the Siwaliks, they’re calling Stegodon insignis or Stegodon ganesa or a hyphenated version of the two: Stegodon insignis-ganesa, which is taxonomic heresy.” 

And here Dr. Jukar was emphatic: “You CANNOT do that with the taxonomic code.”

“And that was Osborn’s fault.”

He was referring to Henry Fairfield Osborn, former professor then curator of the American Museum of Natural History in the late 1800s.

“Osborn [is known to have asserted], ‘I agree with what Falconer said, so I’m going to hyphenate these two words.’  Which created such a mess.  So we have no idea what’s going on there. 

“There’s a lot of work to be done with elephant taxonomy, biogeography and systematics and comparisons between China, the Levant, East Africa and India.”

 

Image of the Levant (Public Domain)

 

Dr. Jukar and other colleagues have also recently published papers on the earliest known fossil of Hexaprotodon, an extinct hippo, from South Asia, and the first record of a Hippaprionine horse (Plesiohipparion huangheense) from the Indian Pliocene.

He is currently working with Dr. Adrian Lister of the Natural History Museum in London to further understand the various proboscidean fossils in the Siwalik collection.

This is important work, but Dr. Jukar pondered its reception to the wider world.

“For a long time paleontologists have been criticized as being mere stamp collectors because we find things and then we name them and then we try to figure out in what larger group they belong to.  But that is the basis of our data.

“Only when I have a comprehensive sense of what the species are, when they lived and where they lived can I start doing these more complicated community-level analyses.

“But because the basic science of naming a fossil might not be very exciting, [as it doesn’t directly impact] human life very much, it doesn’t get a lot of attention. 

“I am definitely interested in the big picture questions of dispersal from Africa into South Asia, about the ecology of these groups, about how communities have changed through time, but I can’t really do a rigorous analysis until I figure out who the [basic] players are in this place.”

Image of Dr. Advait Jukar with a Mammuthus columbi (Columbian mammoth) skull; courtesy of Dr. Jukar

 

References:

  1. Colbert E. (1996). Henry Fairfield Osborn and the Proboscidea. In:  Shoshani J, Tassy P. The Proboscidea : evolution and palaeoecology of elephants and their relatives, Oxford: Oxford University Press, xxii – xxv
  2. Dr. Advait Jukar’s website: https://advaitjukar.weebly.com
  3. Jukar, Advait M., Lyons, S. Kathleen, Uhen, Mark D. (2018.  A cranial correlate of body mass in proboscideansZoological Journal of the Linnean Society, Volume 184, Issue 3, 20 October 2018, Pages 919–931, https://doi.org/10.1093/zoolinnean/zlx108
  4. Jukar, Advait M., Patnaik, Rajeev, Chauhan, Parth R., Li, Hong-Chun, Lin, Jih-Pai (2019). The youngest occurrence of Hexaprotodon Falconer and Cautley, 1836 (Hippopotamidae, Mammalia) from South Asia with a discussion on its extinction, Quaternary International, January 2019, https://doi.org/10.1016/j.quaint.2019.01.005
  5. Jukar, Advait Mahesh, Sun, Boyang, Bernor, Raymond Louis, (2018). The first occurrence of Plesiohipparion huangheense (Qiu, Huang & Guo, 1987) (Equidae, Hipparionini) from the late Pliocene of India,  Bollettino della Società Paleontologica Italiana; 57(2):125-132 · August 2018
  6. Saegusa H. (1996). Stegodontidae: Evolutionary Relationships. In:  Shoshani J, Tassy P. The Proboscidea : evolution and palaeoecology of elephants and their relatives, Oxford: Oxford University Press, xxii – xxv

 

It was a GREAT pleasure and honor speaking with Dr. Advait Jukar.  Many, many thanks for your time, Advait, your help, your fascinating insight and your gorgeous images!! I cannot wait to read your future scientific papers!

 

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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]