Archive for the ‘Pleistocene Mammals’ Category

Hares (Lepus sp.) in Southeastern North America during the Late Pleistocene?

April 17, 2017

Librarians can be a pain in the ass.  On 1 occasion I attempted to check out a book from the Augusta College library.  The librarian told me I needed to purchase an alumni card for the privilege of borrowing a book from my alma mater.  I shelled out $25 for the card, and the same #!#!en librarian still wouldn’t let me take the book home.  Another time I was seeking an old Alabama Journal of Science article.  The authors of the article were dead or in a nursing home so I couldn’t get a copy from them.  The journal posts new issues online but not ones this old.  I contacted a librarian from the Alabama library system and asked her to loan the journal to my nearest library where I could pick it up or at least send me a Xeroxed copy of the article.  I offered to pay for postage and use of the copy machine.  She refused because I was not affiliated with the University of Alabama library system.  My efforts to obtain this article have been stymied for 8 years, but I recently learned a surprising tidbit of information from this article that was referenced in another paper I recently came across.  A tooth identified as comparing favorably to hare was found at Bogue Chitto Creek in Dallas County, Alabama; a site where subfossil remains of late Pleistocene species are occasionally discovered.  Bones of hares have been excavated from 7 sites in Florida that date from the Miocene to the early and mid-Pleistocene, but hares are otherwise unknown from late Pleistocene sites this far south, making this an unique find.

Scientists can’t identify this specimen to a species level based on just this single tooth. Bjorn Kurten, co-author of Pleistocene Mammals of North America, states it is difficult to distinguish between rabbit (Sylvilagus sp.) and hare (Lepus sp.)  teeth, and discerning the difference between hare species based on teeth is even harder, if not impossible.  The tooth may have been from a white-tailed (Lepus townsendii), black-tailed (L. californicus), antelope (L. alleni), or an unknown extinct species of jackrabbit that occupied a small geographic range during the late Pleistocene.  This site is probably too far south for another species of hare–the snowshoe (L. americanus).  It’s also possible the tooth is incorrectly identified and belonged to a true rabbit.  Cottontails have long been abundant all over the south, and they are well represented in the fossil record here.  When paleontologists designate a specimen as comparing favorably (cf), they are not 100% certain of the identification.

Map of Alabama highlighting Dallas County

Bogue Chitto Creek, flows through Dallas County, Alabama.  Many Pleistocene fossil specimens have been found in this creek, including the tooth discussed in this article.

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Present day range map of the  black-tailed jackrabbit.  Western species of hares lived in the southeast during the early to mid-Pleistocene.  Scant evidence suggests they may have occurred in the Black Prairie region of central Alabama during the late Pleistocene as well.

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Black-tailed jackrabbit.  Hares differ from true rabbits.  Their young are born with their eyes open and able to hop about and flee from predators.

Bogue Chitto Creek flows through the Black Prairie region of central Alabama.  The compact clay soils here favor grass over trees, and the Black Prairie region itself extends into neighboring Mississippi and Georgia.  Western hare species prefer large treeless plains, and the predominance of this environment here may explain why a relic population of hares existed in this region during the late Pleistocene.  Other environments in the southeast often climax into forests where western hare species can’t survive.  Lagomorphs (hares, rabbits, and pikas) are susceptible to disease outbreaks, and relic populations of hares in the southeast could have easily succumbed to pestilence.  Before I learned about this tooth, I wondered why there was no evidence of hares in the southeast during the late Pleistocene when arid climates led to a greater prevalence of open environments.  This evidence suggests they may have had a local distribution in some parts of the south then.

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Snowshoe hares turn white in winter and brown in summer.

Unlike their western relatives, snowshoe hares prefer forested environments.  A leg bone of a snowshoe hare was found in Cave ACb-2 in Colbert County, Alabama.  This is the southernmost known occurrence of this species, although this is not far from its present day range.  There is anecdotal evidence snowshoe hares occurred as far south as the Great Smoky Mountains National Park as recently as the early 20th century where they possibly still exist today.  Snowshoe hare remains dating to the late Pleistocene have also been found in 2 other southern states–Arkansas and Kentucky.  They require areas with snowpack on the ground for at least part of the year.

Reference:

Ebersole, Jon; and Sandy Ebersole

“Late Pleistocene Mammals of Alabama: A Comprehensive Faunal Review with 21 Previously Unreported Taxon”

Alabama Museum of Natural History Bulletin 28 December 2011

 

Pine Martens (Martes americana) Lived in Southeastern North America during the Late Pleistocene

April 13, 2017

If humans didn’t colonize North America, I believe the pine marten would have a much wider range than it does today.  Presently,  this small carnivore is confined to boreal and mixed forests in Canada, the northern Rocky Mountains, and upper Maine.  In historical times they also ranged into New England.  During the late Pleistocene pine martens lived at least as far south as northern Alabama, and they probably ranged into the piedmont.  (The fossil record of the southeastern North American piedmont region is poor.  I rely on educated speculation to imagine the faunal composition there.)  Pine marten remains dating to the late Pleistocene have been excavated from Cave ACb-2 in Colbert County, Alabama, as well as 2 sites in Tennessee and 2 in Virginia–far south of their present day range.  Pine martens live in low densities, hunting small mammals and birds on the forest floor and in tree tops.  Unlike their relative, the fisher (M. pennanti), pine martens don’t readily re-establish populations after they’ve been extirpated from a certain area.  Archaeological evidence suggests fishers ranged as far south as north Georgia until European colonization when their range was greatly reduced by increased fur trapping, and they thrive wherever they are re-introduced.  But pine martens struggle to increase their populations when they are re-introduced.

Native Americans killed pine martens using deadfall traps.  A heavy rock was propped up by a stick attached to a piece of meat with a string.  The rock crushed the pine marten pulling at the bait.  Pine martens often fail to replenish their populations after humans begin trapping them in a certain area.  They’ve been able to survive in Canada because this region is more sparsely inhabited by people.  The denser population of humans in the southeast not only trapped out the pine martens but planted agricultural fields and cleared the deep forest habitat they require.  Humans can be just as detrimental to some species of small animals as they are to megafauna populations.

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Pine marten. They are about the size of a small house cat.

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Present day range map of the pine marten.  Most of this range was under glacial ice during the Ice Age.  However, they lived south of the ice sheet at least as far south as Alabama.

Map of Alabama highlighting Colbert County

Fossil evidence of pine marten was found in Cave Acb2 in Colbert County, Alabama.  This is its southernmost known occurrence.

Some scientists speculate evidence of pine martens in north Alabama during the Ice Age suggests the region was covered with boreal spruce forests because this is the type of environment where pine martens occur today.  As I’ve noted in previous blog entries, the Ice Age forest that existed in the upper south then was likely a mixed forest consisting of an extinct temperate species of spruce (Critchfield’s) and hardwoods such as oak, hickory, walnut, elm, etc.  Temperatures were only slightly cooler in this region then than they are today.  I believe humans, not climate change, are the reason for the pine marten’s range reduction.

Reference:

Ebersole, Jon; and Sandy Ebersole

“Late Pleistocene Mammals of Alabama: A Comprehensive Faunal Review with 21 Previously Unreported Species”

Alabama Museum of Natural History Bulletin 28 December 2011

Coyote (Canis latrans) Evolution

March 2, 2017

The coyote is a remarkably adaptable and intelligent animal. The evolutionary history of this species began about 43 million years ago when its ancestors, the caniforms (dogs, bears, weasels, skunks, and raccoons) diverged from the feliforms (cats, hyenas, mongoose, and civets).  The canis genus likely originated in North America over 5 million years ago, having evolved from a primitive wolf-like animal known as eucyon.  Johnston’s coyote (Canis lepophagus) was an early member of the canis genus that lived in North America during the Pliocene from ~5 million years BP-~2 million years BP.  Most paleontologists who study the anatomy of canids believe C. lepophagus was ancestral  to wolves, coyotes, and dogs. Wolves crossed the Bering land bridge and colonized Eurasia, while coyotes stayed in North America.

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Canis lepophagus attempting to scavenge a carcass of a llama defended by a Borophagus, the bone-eating dog.  This illustration depicts a scene that may have occurred during the early Pliocene or late Miocene over 3 million years ago.  Canis lepophagus is thought to be the common ancestor of wolf, dog, and coyote.

At the Rancho La Brea Tar Pits in California coyote bones are the 3rd most common specimens to be excavated here behind dire wolves (Canis dirus) and saber-toothed cats (Smilodon fatalis).  Timber wolf (Canis lupus) skeletal material is present but uncommon.  The abundance of carnivore specimens from this site allows scientists to study changes over time in the anatomy of these species.  Ice Age coyotes from this locality were larger and more powerful than present day coyotes, and they had larger jaws and teeth.  There are probably a couple of reasons for this size disparity.  Pleistocene coyotes hunted larger prey and had a better diet.  They may have hunted juvenile individuals of megafauna species such as horse, bison, camel, and llama; and there was more meat to scavenge.  Moreover, they had to compete with larger carnivores and likely lived in bigger packs.  Dire wolves competitively excluded timber wolves from coyote range.  This benefitted coyotes as well because there was less of an ecological niche overlap between dire wolves and coyotes than there is between timber wolves and coyotes.  Less than 1000 years after the extinction of the Pleistocene megafauna, coyotes evolved to their present day size and stature.

Dire wolves are long gone and timber wolves have been extirpated from most of their former range, but coyote populations are increasing, and they have re-colonized eastern North America within the past century.  Coyotes are 1 of the few carnivores smart enough to avoid poison bait and traps.  Studies show they produce larger litter sizes in response to human hunting pressure.  Though human hunting of coyotes may cause a temporary decrease in their populations, in the long term their populations increase because they begin producing larger litters.  This explains why their populations increase despite being considered a pest that can be hunted year round with no bag limit.

Coyotes in northeastern North America interbred with the last timber wolves in eastern Canada and dogs, and these coyote-wolf-dog hybrids live in large cities and suburbs.  Genetic studies suggest these hybrids are 65% coyote, 25% wolf, and 10% dog.  Characteristics inherited from dogs help them tolerate urban noise, and some have even learned to look both ways before they cross roads.   They’ve adapted well to living on golf courses, city parks, abandoned farmland, vacant lots, cemeteries, and roadside ditches where they have frequent access to roadkill.  I think coyote-wolf-dog hybrids have colonized southeastern North America as well.  I’ve seen some that look like western coyotes, and others that resemble wolves.  More genetic studies of southeastern coyotes may confirm my hypothesis.

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Coyote-wolf hybrids have colonized northeastern North America.  I hypothesize some coyotes in southeastern North America have also bred with wolves and dogs.

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These photos help distinguish between coyote-wolf hybrids and pure bred coyotes.  I’ve seen canids in Georgia that resemble both.

Reference:

Meachen, Julie; and Joshua Samuels

“Evolution in Coyote (Canis latrans) in Response to Megafaunal Extinctions”

PNAS 2012

The Unknown Mating Habits of Saber-toothed Cats (Smilodon fatalis)

February 5, 2017

The average male saber-toothed cat was only slightly larger in overall body size than a female saber-tooth, but they had significantly larger mandibles and upper canines (the fangs).  This is in contrast to most species of cats today.  Most male cats and especially lions are much larger than the average female of their species.  The mating habits of Smilodon are completely unknown, and we can only speculate about them based on the knowledge that they differed in size above the neck, but not much elsewhere aside from the sexual organs.

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Smilodon had low sexual dimorphism in body size, but males had significantly larger mandibles and upper canines.  Did males share their larger kills with females as way to attract them?

I believe saber-tooth mating habits may have been notably different from those of all extant species of cats.  The saber-toothed and the scimitar-toothed cats ( Dinobastis or Homotherium ) belonged to an extinct subfamily of cats known as the Machairodontinae.  The Machariodontinae diverged from all other cats an estimated 13 million years ago, very early in cat evolution, and they have no close living relatives today.  They were more closely related to other carnivores than modern day species of cats are.  Perhaps they lived in matriarchal societies like the spotted hyena ( Crocuta crocuta ), another species that shows low sexual dimorphism (the females are actually slightly larger than the males).  Or maybe, as in the wolf ( Canis lupus only the dominant male and female of the pack were allowed to mate.  However, scientists disagree over whether saber-toothed cats were social or solitary animals.

Some scientists argue evidence from the La Brea Tar Pits of severely injured saber-tooths that survived traumatic debilitating injuries suggests they must have lived in groups.  But others believe that even a severely injured saber-tooth could have lived for a long time by scavenging.  A saber-tooth in a bad mood due to pain could have easily intimidated smaller predators from their kills. Moreover, their small braincases also indicates they didn’t live in groups.  I suspect they were solitary cats, though mothers probably hunted cooperatively with nearly grown cubs when she was training them how to hunt.

The males were able to bring down larger prey than the females because their bites, aided by the larger jaws and fangs, were deadlier.  Perhaps this was an element of their mating system.  Females that came to scavenge the male-killed prey were tolerated by the males, and the heavy meal caused them to go into instant heat.  Maybe females followed the most successful hunting males with the frequent nutritious meals triggering more frequent ovulation.

I’ve always been fascinated over how recently this strange exotic animal became extinct–only ~10,000 years ago.    It’s frustrating not to be able to know more about how it lived.  Relying on guesswork is just not as satisfying as knowing.

References:

Christiansen, P.; and John Harris

“Variation in Craniomandibular Morphology and Sexual Dimorphism in Pantherines and the Sabercat Smilodon fatalis

Plos One Oct 2012

Meachen-Samuels; J.A.; W.J. Binder

“Sexual Dimorphism and Ontogenetic Growth in the American Lion and Sabertoothed Cat from Rancho La Brea”

Journal of Zoology 2010

Mammoths and Mastodons were Year Round Residents of the Ohio River Valley during the Late Pleistocene

January 27, 2017

The bones and teeth of an extinct animal provide scientists with information about the life history of that particular individual.  Recently, 2 scientists analyzed the chemistry of 8 mammoth ( Mammuthus columbi ) and 4 mastodon ( Mammut americana ) teeth collected from Hamilton County, Ohio and Bullitt and Gallatin Counties, Kentucky–in other words, the Cincinnati area.  They determined 11 of these animals spent their entire lives in what was to become the Cincinnati area.  They also learned 7 of the mammoths ate different plant foods than the mastodons, but the diet from 1 individual mammoth did overlap with mastodon diets.

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Hamilton County, Ohio

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Mammoths and Mastodons lived year round in the Ohio River Valley.  They were 2 completely different species of proboscidean.

Scientists are able to determine where an animal lived during its lifetime by measuring the ratio of strontium and strontium isotopes in their teeth.  Strontium leaches from local bedrock into the water supply, and animals absorb the strontium into their bones when they drink.  Different geographical regions exhibit different ratios of strontium isotopes, so it’s possible to figure out where an animal spent time during its life.  A previous study of mastodon and mammoth teeth collected from Florida determined mastodons there migrated back and forth from central Florida to central Georgia, while Florida mammoths did not migrate.  But this study suggests most of the mastodons that lived in the Ohio River valley did not migrate.  However, there were exceptions.  The strontium ratio from 1 specimen indicated this individual wandered north from either north Georgia or southern Tennessee to the Ohio River valley.  The authors of this study estimated this could have been accomplished in as little as 5 days based on how fast modern elephants can travel.

The bone chemistry tells us mammoths mostly ate grass, while mastodons ate plants that grew in forested environments.  But again there was 1 exception–1 mammoth that fed upon forest vegetation.  During the Last Glacial Maximum much of this region was a cool arid steppe environment.  After the nearby ice sheet retreated, the steppe was transformed into an open spruce parkland, then eventually an oak and hardwood dominated forest.  This mammoth apparently adapted to the latter changes.

Mammoths and mastodons had no need to migrate away from the Ohio River.  The rich floodplain habitat and numerous mineral licks provided enough nutrition to support year round populations of both.

Reference:

Baumann, Eric; and Brook Crowley

“Stable Isotopes Reveal Ecological Differences among now Extinct Proboscideans from the Cincinnati Region, USA”

Boreas 2015

Donald Grayson’s Disingenous Case Against Overkill

January 23, 2017

I almost chose not to read Donald Grayson’s most recent book, Giant Sloths and Sabertooth Cats: Extinct Mammals and the Archaeology of the Ice Age Great Basin.  Grayson is a long time skeptic of the hypothesis that man overhunted Pleistocene megafauna to extinction, and he has authored and co-authored a number of papers explaining his position.  In my opinion overhunting by man is the only explanation for the extinction of Pleistocene megafauna that makes sense.  I’ve read his papers and consider his arguments highly illogical and unfair.  But I did purchase his book because I try to absorb all the knowledge I can about my favorite subject–the late Pleistocene ecology of North America.  I don’t have to agree with an author about everything to enjoy their work.  I saw his chapter on extinction was short, just a small portion of the book, and I assumed he would simply rehash his tired old case against overkill.  However, I was shocked at the way he misrepresented the results of a paleoecological study.  He implies the results of this study support climate change models of Pleistocene megafauna extinctions.  In fact this paper specifically states the opposite.

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Donald Grayson’s newest book.  His chapter on extinction is marred by gross deception.

Don Grayson, photo by Mary Levin, UW Photography, 2011

Donald Grayson is an archaeology professor at the University of Washington.  

On page 287 of the above book, Grayson wrote “…paleoecologist Jacquelyn Gill and her colleagues took a close look at tiny bits of Ice Age history extracted from 3 lakes and marshes in New York and Indiana.”    Grayson fails to mention the data from the lakes in New York was gathered in a study led by Guy Robinson (1).  (I will number the studies mentioned in this blog post and list them below.)  Guy Robinson’s team took samples of sediment from some New York lakes and marshes, radiocarbon dated different chronological layers of it, and measured and analyzed the volume of pollen, charcoal, and dung fungus spores in each layer.  The volume of dung fungus spores is used as a proxy to estimate populations of megafauna.  They determined local extinctions of megafauna were staggered throughout 2 thousand years, and they concluded this ruled out climate change as a cause of the megafauna extinctions.  If climate change caused the extinctions, they believed the extinctions at these different sites would be simultaneous.  Instead the local extinctions occurred at different times at different sites, and it appears as if nomadic humans were overhunting megafauna at 1 site, then moving on to another, though these local extinctions occurred shortly before the Clovis cultural era.  Increases in charcoal while megafauna populations were declining is additional evidence of probable human impact.  Jacquelyn Gill, the lead author of the paper (2) Grayson referred to in his book,  details a similar study of an Indiana lake.  The authors of this study also found that megafauna populations declined before the Clovis cultural era but also before climate caused changes in the local plant composition.  They believe human impacts are still a “plausible” cause of megafauna extinction, and they wrote “our data thus rule out the hypothesis that (i) climate-forced changes in vegetation drove the megafaunal decline, and (ii) no-analogue plant communities were created by megaherbivory.”  Megafauna became locally extinct here before the pollen evidence suggests changes in the plant community occurred.  They believe humans can still be implicated in megafauna extinctions at this site because increases in charcoal indicated humans were setting fire to the landscape during the period of megafauna decline, and there is evidence of human butchery of mammoths in southeastern Wisconsin during this same time period, suggesting that humans were likely hunting them at this location as well.

It is very dishonest of Grayson to imply these studies support climate-change models of extinction when the authors of these studies specifically state the opposite.  He should have at least informed his readers of their conclusions and explain why he has a different interpretation of the data.  The authors of these studies do note that megafauna decline occurred during a warm phase of climate, but they believe, if climate played a role, it had to be some other mechanism besides climate-driven changes in the environment.  The accepted logic behind climate change models of extinction is that changes in climate decreased the plant foods necessary to maintain viable populations of megaherbivores.  These studies show this is not the case.  Moreover, fossil coprolites show the plants Ice Age megafauna ate are still common on the landscape today, and isotopic studies indicate they were not picky feeders, but instead ate a wide variety of foods.  I think the warm climate phase provided a greater variety of edible plant foods for humans, thus increasing human populations which in turn was detrimental to megafauna.

There are many regions in North and South America where the environment did not change substantially during the most recent glacial-interglacial transition including the pampas of Argentina, southern California, and southeastern North America, especially Florida and the coastal plain.  Climate change models of extinction just don’t make sense in these regions.  Most species of megafauna enjoyed continent wide distributions, and they endured 30 glacial-interglacial transitions over the past 2 million years without suffering extinctions, yet they did become extinct about the same time man appears in the archaeological record.  This can’t be coincidence.

Grayson focuses his arguments against overkill on the Clovis blitzkrieg model of extinction.  This model proposes humans rapidly overhunted megafauna to extinction within the 500 year period of the Clovis cultural era.  There is another model of overkill: humans were responsible for overhunting megafauna, but it took place over several thousand years and also involved a change in fire regime and other human impacts.  This is known as the protracted overkill model and it is the one I favor.  Grayson ignores this model.  He doesn’t acknowledge the likelihood that pre-Clovis humans impacted megafauna populations.  I’m sure he would point out the lack of archaeological evidence for this.  His main objection to the blitzkrieg model is also a lack of archaeological evidence…he believes there are not enough kill sites (direct evidence that humans killed a beast) to justify the overkill hypothesis.  This objection is unreasonable.  99.999…etc% of the individuals of a species that ever lived left no fossil evidence at all.  It is ridiculous to expect to find fossil evidence of the last individuals of a species that just happened to be killed by men.  Moreover, Grayson never quantifies how many kill sites archaeologists would need to find before he would be convinced humans were responsible for the demise of the megafauna.  He needs to put an exact number on it or cease his objections.  Despite the odds against it, there are numerous kill sites and evidence of human-butchered megafauna bones.  Grayson dismisses over half of them, sometimes unfairly, though sometimes his skepticism is warranted.

Grayson falsely claims the overkill hypothesis has “little going for it” and he sounds annoyed on pages 279 and 280 of his book when he laments its acceptance in the popular media.  He might be annoyed because in recent years the  overwhelming number of paleoecological and statistical studies suggest man at least played some role in megafauna extinctions.  Grayson glosses over a statistical study led by G.W. Prescott (3) that determined both man and climate played a role in the end Pleistocene extinctions.  But 2 recent studies of worldwide extinction chronology (4) (5) determined extinctions are more closely tied to human expansion than climate change.  In the study led by C. Sandom they note extinctions have been severe in climatically stable regions.  They write: “Human arrival was a necessary factor for extinctions, whereas climate variation was a contributory one, enhancing regionally the effects of anthropogenic impacts on additive rather than synergistic ways.”

References:

(1) Robinson, G.S. ; L.P Burney and D.A. Burney

“Landscape Paleoecology and Megafaunal Extinction in Southeastern New York”

Ecological Monographs 2005

(2) Gill, J.L.; J.W. Williams, S.T. Jackson, K.B. Lininger, and G.S. Robinson

“Pleistocene Megafauna Collapse, Novel Plant Communities, and Enhanced Fire Regimes in North America

Science 2009

(3) Prescott, G.W. et. al.

“Quantitative Global Analysis of the Role of Climate and People in Explaining Late Quaternary Extinctions”

PNAS 2012

(4) Aravjo, Bernardo et. al.

“Bigger Kill than Chill: the Uneven Roles of Humans and Climate on Late Quaternary Megafaunal Extinctions”

Quaternary International 2015

(5) Sandom, C. et. al.

“Global Late Quaternary Megafauna Extinctions Linked to Humans, not Climate Change”

Proceedings of the Royal Society 2014

http://rspb.royalsocietypublishing.org/content/281/1787/20133254

 

A Recent Study of Pleistocene Armadillo DNA Yields 2 Surprising Results

January 19, 2017

An extinct species of armadillo ( Dasypus bellus ) ranged throughout southeastern North America during the Pleistocene.  (A much larger species, Holmesima septentrionalis, was restricted to Florida and the lower coastal plain.)  Scientists have described D. bellus , known by the common name of beautiful armadillo, as being remarkably similar to the extant 9-banded armadillo ( D. novemcinctus ).  The most notable difference between the 2 species is size–the beautiful armadillo was twice the size on average as the 9-banded armadillo.  The latter species began to expand its range into southeastern North America from Mexico within the last 150 years, and today is very common and on the increase in the region.  In a previous blog entry I hypothesized the 9-banded armadillo was a dwarf mutation of the beautiful armadillo, and it was currently recolonizing former parts of its range.  (See: https://markgelbart.wordpress.com/2012/08/19/is-the-9-banded-armadillo-dasypus-novemcinctus-a-dwarf-mutation-of-the-pleistocene-species-dasypus-bellus/ ) However, scientists were recently able to extract DNA from 2 Pleistocene-aged armadillo specimens, and they determined the history of the 2 Dasypus species is more complicated and even more interesting than previously thought.

An armadillo I saw at Scull Shoals State Park, Georgia.  2 species of similar armadillos occupied southeastern North America during the Pleistocene including this 1.

Scientists extracted DNA from an armadillo specimen found in Brynjulfson Cave, Missouri and from another specimen excavated from Medford Cave, Florida.  (They tried many other specimens but these were the only 2 that still yielded viable DNA.)  They determined the DNA of the Missouri specimen was distinct enough from modern 9-banded armadillo DNA to be considered a distinct species.  So much to my surprise, the beautiful armadillo is not the same species as the 9-banded armadillo.  But the Florida specimen held an even bigger surprise…it was a 9-banded armadillo and it dated to over 10,000 years ago.  This means both beautiful armadillos and 9-banded armadillos lived in southeastern North America during the Pleistocene.  The former went extinct while the latter was temporarily extirpated from the region but has just recently made a comeback.

The scientist who originally described the specimen from Florida noted its similarity to the 9-banded armadillo but chose to identify it as a beautiful armadillo because that was the species thought to occur there during the Pleistocene.  This individual was as large as a beautiful armadillo, showing that size alone is not enough to diagnose correct species identification.  Unfortunately, most subfossil specimens no longer contain DNA due to permineralization or decay.  All the specimens labeled ” D. bellus ” in the scientific literature should be re-labeled as ” D. species ” until scientists make a more detailed anatomical analysis of the genus, so that these 2 species can be better distinguished.

So why did the 2 species disappear from southeastern North America near the end of the Pleistocene?  Cold arid climate cycles probably caused range reductions and local extinctions, but armadillos likely re-expanded during warmer wetter climate phases.  Today, 9-banded armadillos may use manmade roads to facilitate their range expansion because it’s less strenuous to travel along cleared roadsides (though dangerous because highways are littered with armadillo corpses). During the Pleistocene armadillos probably followed trails trampled clear by herds of megafauna. This facilitated range expansion during favorable climate cycles.  The extinction of the megafauna may have played a role in the demise of armadillos that could no longer expand their range after climate deterioration caused extirpations. This isn’t a completely adequate explanation–Florida never got too cold and dry for armadillos.  The authors of the below referenced study suggest frequent manmade fires may have been detrimental to armadillos.  Native Americans set fire to the woods annually.  Modern day fire suppression may be another reason 9-banded armadillos have been able to recolonize former territory.

Reference:

Shapiro, Beth; Russell Graham, and Brandon Letts

“A Revised Evolutionary History of Armadillos (Dasypus) in North America Based on Ancient Mitochondrial DNA”

Boreas August 2014

 

 

 

 

Pleistocene Mammals of the Levant

January 14, 2017

Long before the stories in the bible supposedly took place, the Levant was a beautiful wilderness sparsely populated by humans.  The Levant is the region encompassing the modern day boundaries of Israel, Lebanon, Syria, Jordan, and Iraq.  For millions of years climatic fluctuations have caused a waxing and waning of 2 different types of environments here–Mediterranean evergreen oak woodlands and Irano-Turanan steppe consisting of deciduous oak trees and grassy understories.  Habitat for both forest species and grassland fauna has been available during every climatic stage.  The region is also a gateway between Eurasia and Africa, so animals from 3 continents converge here, making it rich in diversity.  African species such as elephants, giraffes, rhinos, hippos, gazelles, hartebeest, warthog, macaque, hyena, lion, leopard, cheetah, and Cape Hunting dog formerly lived side by side with Eurasian species including aurochs, bison, horse, ass, camel, deer, wild boar, ibex, wolf, and brown bear.

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Map of the Levant.

The fossil record suggests the fallow deer ( Dama dama ) was the most common large herbivore in the Levant for over 2 million years.  This species prefers fairly dense woodlands, so their abundance in the fossil record surprises me because I always think of this region as arid.  However, during Ice Ages, the climate in the Levant was cooler and rainier than it is today, though drier climate phases did occur cyclically.  The extinct giant deer ( Megaloceros giganteus ) and elk ( Cervus sp. ), known as red deer in Europe, also made the Levant their home.  The wild ibex ( Capra aegargus ), ancestor of the domestic goat ( C. hircus ), was common on rocky hillsides; gazelles, hartebeest, and an extinct species of warthog roamed the grassy plains.

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Fallow deer.

During warmer climate cycles hippos inhabited Lake Kinnaret.  Long chains of lakes often existed along the Jordan River, and during some climatic stages Lake Kinnaret joined the extinct Lake Amora and the Dead Sea to become 1 giant primeval lake known as Lake Lisan.  Oddly enough, geologists believe Lake Lisan was a freshwater lake in the part that covered the current site of Lake Kinnaret, while the rest of the lake was salty.

A primitive genera of elephants known as stegodon became extinct in Africa about 1 million years ago, but they still lived in the Levant for hundreds of thousands of years past their African extinction.  Stegodon survived until the end of the Pleistocene in southeastern Asia.  Two species of elephants roamed the Levant during the Late Pleistocene–the steppe mammoth ( Mammuthus trogontherii ) and the straight-tusked elephant ( Paleoloxodon antiquus ).  The former evolved into the woolly mammoth during a later Ice Age.  Straight-tusked elephants were a temperate species that couldn’t survive the climate deterioration of the last Ice Age in most of Eurasia.  However, the Levant probably provided a refuge for this species then.  I hypothesize humans overhunted straight-tusked elephants to extinction in their final refugia.  And I believe the same fate befell the temperate species of rhino ( Stephanorhinus hemiotoechus ) that occurred throughout Eurasia.  The Levant likely served as a refuge for these 2 species of megafauna during previous glacials, but human populations and/or hunting skills increased enough to permanently eliminate these slow breeding animals sometime within the timespan of the most recent Ice Age.

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Stegodon.

The lion ( Panthera leo ) that lived in the Levant was the same subspecies as the Asiatic lion found today in 1 small area of India–the Gir Forest.  This big cat survived in remote regions of the Levant until the 19th century.  There is still a small population of leopards in the Levant.  Two species of wolves ranged through the Levant–the timber wolf ( Canis lupus ) and the Egyptian wolf ( C. lupaster ).   Though the latter species occasionally interbreeds with golden jackals ( C. aureus ), a genetic study determined they are more closely related to C. lupusA single specimen of Cape Hunting dog ( Lycaon pictus ) was excavated from Hayonim Cave, Israel.  The paper written about this site incorrectly states this as the only fossil material of Cape Hunting dog ever found outside Africa, but fossils of closely related species have been discovered in Alaska and Texas.

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Asiatic lions in Gir Forest–the same subspecies lived in the Levant until the 19th century.

Pleistocene megafauna suffered fewer extinctions in the Levant than in the Americas.  Wildlife there co-occurred for a longer time with low populations of primitive humans and had time to evolve better avoidance strategies.  Moreover, many Levant species that did become extinct in the wild still live on as domesticated descendents.  Nevertheless, most of the megafauna species were extirpated from the Levant by the 20th century.

References:

Marder, Ofer; et. al.

“Mammal Remains of Rantis Cave, Israel and mid to late Pleistocene Paleoenvironment and Subsistence in the Levant”

Journal of Quaternary Science 2011

Stimer, Mary; and Ofer Bar-Yozef

“The Fauna of Hayonim Cave, Israel: A 200,000 Year Record of Paleolithic”

American School of Prehistoric Research 48 2009

The Super Squirrel

January 3, 2017

The eastern gray squirrel ( Sciurus carolinensis ) outlasted many of the magnificent extinct species of Pleistocene megafauna because they are well adapted to survive in environments modified by man.  They are just as much at home in suburbs, city parks, and 2nd growth forest of the countryside as they are in the middle of a pristine wilderness.  Unlike western gray squirrels ( S. griseus ), they are not shy around man and will nest in backyards or even attics.  They are nimble squirrels, able to jump from tree top to tree top in the young dense forests that replace abandoned agricultural lands.  And they have a unique way of spreading their populations.  Every September, juvenile eastern gray squirrels begin to expand their range and forage for acorns and nuts.  After they have spent enough time  burying acorns in a certain area, they establish an home range there.  This process is known as the “September shuffle.”  During colonial times when there were still vast tracks of timber, this September shuffle could seem like a massive migration, especially during years of poor mast production following a year of heavy mast production that increased squirrel numbers. (See: https://markgelbart.wordpress.com/2012/08/24/squirrel-migrations/ )

Eastern gray squirrels thrive everywhere they’ve been introduced–England, Ireland, Italy, South Africa, western Canada, and Australia (where they were eventually eradicated by man).  Much to the consternation of English naturalists, they have almost completely displaced native European red squirrels ( S. vulgaris ) on the British Isles.  There are several possible reasons for this displacement.  Eastern grays are carriers of a virus that may be fatal to European red squirrels.  They may also disrupt red squirrel mating and outcompete them for food, and they are simply better adapted to living adjacent to people.  But the most compelling ecological explanation involves comparing European and American Ice Ages.

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The American eastern gray squirrel in Brandon Hill Park, Bristol, England.  Gray squirrels are better adapted to living in deciduous woodlands than native red squirrels and they are displacing them.

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The Eurasian red squirrel is being displaced by introduced American gray squirrels in Great Britain and Italy.

In Europe glaciers covered a greater percentage of territory than they did in North America.  Most of the unglaciated region consisted of grassy or shrubby mammoth steppe with pockets of spruce and pine growing in moist protected areas.  Some southerly lowlands supported more extensive conifer forests.  The deciduous oak forests that dominate most of Europe today were restricted to narrow strips along the Mediterranean coast.  Because glacial stages were 5-10 times longer than interglacials, European red squirrels became better adapted to live in conifer forests.  However, in North America, even during the severest stadials, there were always extensive oak and oak/pine forests that supported large populations of gray squirrels.  Eastern grays evolved the ability to digest acorns better than red squirrels can.  Although eastern grays are not native to Europe, they are a better fit for the interglacial oak forests that exist there today.  Ecological displacement of 1 species by another has occurred thousands of times during earth’s history.  People may object to the displacement of European reds by eastern grays because man played a role in the introduction of the latter, but it is not unnatural or detrimental to the overall ecosystem in this case.

Eastern grays along with California ground squirrels, introduced fox squirrels ( S. niger ), and turkeys are displacing and outcompeting western gray squirrels on the Pacific coast of North America.  All seem to be better adapted to anthropogenic environments.  Western grays are now restricted to deep wilderness preserves.  Introduced eastern gray squirrels are also displacing American red squirrels ( Tamiasciurus hudsonicus ) in British Columbia.

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The American red squirrel is being displaced by gray squirrels in some parts of its range as well.

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Taxidermic mounts of eastern and western gray squirrels. The western is larger with a bushier tail.  Introduced Eastern gray squirrels and fox squirrels, as well as native California ground squirrels and turkeys are outcompeting western grays in suburban areas of California heavily modified by man.

Eastern gray squirrels have co-existed with fox squirrels for hundreds of thousands of years but are more common in many areas, including Richmond County, Georgia where I live.  Eastern grays quickly recolonize agricultural land replaced by dense 2nd growth forest, while fox squirrels prefer mature forests with widely spaced trees.  Because most of southeastern North America was clear cut between 1865-1945, gray squirrels have been quicker to return and spread throughout their range.  Without human introduction fox squirrels may never return to formerly clear cut land.

Eastern gray squirrels boldly live next to people and their unique September shuffle makes them a super squirrel, able to expand their populations and survive where other squirrels can’t.

Reference:

Bruemmer, Corrie; Peter Lurz, Karl Larsen, and John Gurnell

“Impact and Management of Alien Eastern Gray Squirrel in Great Britain and Italy: Lessons from British Columbia”

Proceedings of the Conference on the Biology and Management of Species and Habitats at Risk 1999

 

 

Black Bear (Ursus americanus) Diversity during the Pleistocene

December 19, 2016

Fossil evidence suggests North American black bears evolved from an Holarctic population of bears about 3 million years ago.  The founding preceding species is known as Ursus abstrusus in North America and Ursus minimus in Eurasia but they were likely the same animal.  Eurasian black bears ( Ursus thibetanus ) diverged from North American black bears during some climate phase when the ancestral populations became geographically isolated.  Before this divergence moderate climate allowed forested conditions to exist across the Bering land bridge.  But deteriorating climate transformed the land bridge to tundra when it wasn’t submerged under the Bering Sea.

Genetic evidence indicates western populations of North American black bears diverged from their eastern counterparts about 1.8 million years ago.  This corresponds with the beginning of the Pleistocene.  Although weak Ice Ages occurred during the preceding Pliocene, they became much more severe at the onset of the Pleistocene.  Glaciers covered most of Canada and the upper elevations of the Rocky Mountains, blocking gene flow between eastern and western populations of black bears.  Some mixing occurred (and is presently occurring) during interglacials, but because glacial climate phases are 5-10 times longer than most interglacials, isolation between eastern and western populations has  been the norm.

An ancient isolated population lives along the coast and islands of British Columbia.  Glaciers covered most of British Columbia during the Last Glacial Maximum but a strip of land along the coast, including the now submerged continental shelf, hosted a temperate rain forest with a population of black bears.  These forests were probably snow-covered for much of the year, an environmental condition that may have selected for white bears.  The white color may also help improve success hunting for salmon.  The lighter color is harder for the fish to see during the day. Today, this region is home to the highest incidence of white-coated black bears, variously known as Kermode or spirit bears.  On mainland British Columbia 1 in 40 black bears have white coats, while on some of the British Columbian islands 1 in 8 have white coasts.  (They are not albino.)  White bears in most black bear populations are extremely rare in other regions, though when the species was more common there were occasional individuals with white coats.  A white-coated bear was killed in a 1760 ring hunt (See: https://markgelbart.wordpress.com/2014/07/27/the-pennsylvania-mammal-holocaust-of-1760-a-rare-record-of-an-old-fashioned-varmint-drive/ )  Black bears may also be cinnamon, blonde, or even blue.

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The Kermode or Spirit bear–a white color phase of the black bear most common along the coast of British Columbia.  It’s an Ice Age relic population.

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Blue color phase cubs of a black bear mother.

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Cinnamon color phase mother with black and blonde (?) phase cubs.

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Blonde black bear.

During the Pleistocene before humans reduced black bear populations, many grew as large as grizzlies, and they had a much greater genetic diversity.  However, they competed with giant short-faced bears ( Arctodus simus ) and Florida spectacled bears ( Tremarctos floridanus ).  In open environments I think the former may have excluded black bears much in the same way grizzlies kept black bears from ranging into California valleys.  Spectacled bears co-existed with black bears in the same forested habitats for over a million years, and the environment in many places was rich enough to support both species.  Black bears were more adaptable than both of these now extinct species.  I hypothesize that unlike giant short-faced bears, they learned to fear man.  The ability to hibernate during cold weather also made them more widespread and successful than spectacled bears which were probably limited to regions with warmer climate.

Reference:

Marshall, H.D. and K. Ritbad

“Genetic Divergence and Differentiation of Kermode Bear Populations”

Molecular Ecology 2002

Wooding, Stephen and R. K. Ward

“Phylogeography and Pleistocene Evolution in North American Black Bear”

Molecular Biology and Evolution  1999