The Relationship Between

Geographic Barriers and Divergence

Katie CarilloLauren DavisCandice EspinozaAntonio Marquez

University of New Mexico 2012

Table of Contents

Introduction to Barrier Driven Divergence 2

Activity 1: Related Squirrel Subspecies in 5

the Grand Canyon Area

The Grand Canyon as a Geographic Barrier 7

Types of Barriers 9

Crossing Barriers 11

Gene Flow 12

Modes of Speciation 13

Potential Effects of a Man-made Barrier on 15

Wildlife

Activity 2: Big Horned Sheep and Pygmy 17

Owls

Further Investigation 19

1

Introduction to Barrier Driven Divergence

There are two populations of Abert’s squirrels (Sciurus aberti) that currently live in the area surrounding the Grand Canyon in Arizona. The squirrel populations share common ancestors but have long been separated by the Grand Canyon, and divergence has begun to occur producing two different subspecies, Sciurus aberti kaibabensis and Sciurus aberti ab-erti.

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Both subspecies have tasseled ears and live in Ponderosa forests feeding on Ponderosa pine cones, other nuts, seeds, and fruits. There are clear phenotypic differences between the two; Sciurus aberti kaibabensis on the North rim of the Grand Canyon have black bellies and light tails while individuals belonging to Sciurus aberti aberti on the South rim have white bellies and dark tails. Visual differences between the two subspecies are so great, that scientists previously assumed that they were two separate species. Mo-lecular (DNA) evidence made it clear that the two squirrel populations were more closely related.

…What caused them to diverge?

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Figure 1. Notice S. a. aberti (left) has a white belly while S.a. kaibabensis (right) has a dark belly.

Figure 2. Notice S. a. aberti (left) has a dark tail while S.a. kaibabensis (right) has a white tail.

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1. Go to the Arctos database (http://arctos.database.museum/home.cfm) and click on the Museum of Southwestern Biology link.

2. Type in Sciurus aberti kaibabensis �QV�\PM�1LMV\QÅKI\QWV�IVL�Taxonomy box.

3. Type Arizona in the Locality box.

4. When the specimen list pops up, click on the button at the top that says Berkeley Mapper & Rangemap. It will open a new page and show a map of this subspecies range. What is the range of this species?

5. Go back to Arctos and choose specimen 191256 from the Grand Canyon National Park.

6. When was the specimen collected?

7. What body parts were retained?

8. What sex was the specimen?

9. Copy the coordinates at which the specimen was collected and paste them in Google Earth http://www.google.com/earth/index.html

Activity 1: Related Squirrel Subspecies in the Grand Canyon Area

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10. Go to GenBank http://www.ncbi.nlm.nih.gov/genbank/. Select Taxonomy from the drop box and type Sciurus aberti kaibabensis. In the right hand corner there is a box which has nucleotide and protein sequences. Click on the number two in the box next to nucleotide. Choose Mitochondrial cytochrome b. Go to the bottom and look at the nucleotide sequence. Leave this page open.

11. Go back to Arctos and type in Sciurus aberti aberti.

12. Choose specimen 191249.

13. Answer questions 4, and 6-10.

14. Compare the DNA sequences for Mitochondrial cytochrome b for the two subspecies. Are there differences? Hint: Look very carefully at segment starting with nucleotide 101, and 171. (note: nucleotide 104 and 176 are not the same in the two subspecies). Are there other nucleotide differences?

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The Grand Canyon as a Geographic Barrier

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The Grand Canyon is 227 miles long, 18 miles wide, and 6000 feet deep. It was carved out by the Colorado River over 17 million years ago and Sciurus aberti aberti has long been a resident of the Grand Canyon area.

Sciurus aberti kaibabensis lived in Ponderosa forests on the Kaibab plateau when the climate was warmer and the Pon-derosa’s were limited in range. The squirrels were isolated from other populations of Abert’s squirrels due to discontinuation of Ponderosa pine habitat, and divergence began. When the climate started to cool again the range of Sci-urus aberti kaibabensis expanded back down to the edge of the Grand Canyon with the Ponderosa forests.

...Have the squirrels been separated for 17 million years?

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Types of Barriers

The Grand Canyon is an example of a geographic barrier that caused the divergence of Abert’s squirrels into two subspecies. However, canyons are not the only forms of geographic barri-ers. Other naturally formed barriers include:

Mountains Rivers Oceans Deserts

Human made structures can also act as barriers, such as:

Walls Highways Dams

.... Can you think of others?

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10

Crossing Barriers

A species’ habitat and level of mobility determines what barriers that species can cross. A mountain range may cre-I\M�I�JIZZQMZ�NWZ�I�UW][M��J]\�\PMZM�IZM�OMM[M�\PI\�KIV�Æa�W^MZ�\PM�0QUITIaI[��)�KIUMT�KIV�KZW[[�I�LM[MZ\��I�Å[P�KIVVW\��<PM�oceans create a barrier for most terrestrial birds and mammals, but a whale can swim from one ocean to another. Other factors \PI\�UIa�KWV\ZQJ]\M�\W�UWJQTQ\a�QVKT]LM"�IJQTQ\a�QVIJQTQ\a�\W�Æa�high off the ground or long distances, and resource availabil-ity. Biologists need to know the natural history of a species to understand the impact of barriers on divergence.

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Gene Flow

� /MVM�ÆW_�Q[�\PM�\ZIV[NMZ�WN �OMVM[�NZWU�WVM�XWX]TI\QWV�\W�another population. An important factor that affects the rate of OMVM�ÆW_�JM\_MMV�\_W�XWX]TI\QWV[�Q[�UWJQTQ\a� Greater mobility gives rise to increased migratory poten-tial, and migration into or out of a population may be respon-sible for changes in allele frequency and/or the addition of new genetic variants.� 5IQV\IQVML�OMVM�ÆW_�JM\_MMV�\_W�XWX]TI\QWV[�ZML]KM[�divergence between the groups by admixing the two gene pools. 7V�\PM�W\PMZ�PIVL��IJ[MVKM�WN �OMVM�ÆW_�JM\_MMV�\_W�XWX]TI-tions can increase genetic divergence by allowing them to change independently. Ultimately divergence can lead to speciation.

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Modes of Speciation

There are four primary modes of speciation: allopatric speciation, parapatric speciation, peripatric speciation, and sympatric speciation.

Allopatric speciation occurs when biological popula-tions of the same species are physically isolated by a geographic barrier.

Parapatric speciation takes place when the ranges of two same species popula-tions are immediately ad-jacent to each other often forming a contact zone.

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Peripatric speciation oc-curs when a small popula-tion is isolated at the edge of a larger population of the same species.

Sympatric speciation is the process through which new species evolve from a single ancestral population while occupying the same geographic area.

...Divergence could possibly lead to what mode of speciation between Sciurus aberti kaibabensis and Sciurus aberti aberti squirrels?

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Potential Effects of a Man-made Barrier on Wildlife

There is currently a 1125-km security fence under con-struction in United States spanning more than one-third of the U.S.–Mexico border. The fence reaches heights greater than 4 meters and is the cause of large vegetation clearings. The com-bination of the height of the wall and the large spans of cleared land could greatly impact the movement of wildlife species living in these habitats. Animal movements determine distribu-\QWV��IJ]VLIVKM��IVL�OMVM�ÆW_���5IVa�IVQUIT�XWX]TI\QWV[�_QTT�be affected by the construction of U.S. border wall; two of the animals that may be affected are the pygmy-owls and bighorn sheep.

Pygmy-Owls are small nocturnal owls that have an aver-IOM�ÆQOP\�PMQOP\�WN �WVTa�����UM\MZ[�IJW^M�OZW]VL��\PMZMNWZM�\PMa�_QTT�VW\�JM�IJTM�\W�Æa�W^MZ�\PM�_ITT�VWZ�KZW[[�TIZOM�^MOM\I\QWV�gaps.

Bighorn Sheep often form small and fragmented popula-tions making connectivity between the population important. There are currently nine populations of bighorn sheep in north-western Sonora that are linked by dispersal with those in border-ing Arizona. Populations on either side of the wall may become isolated if corridors are disrupted by impermeable fencing.

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Activity 2: Big Horned Sheep and Pygmy Owls

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1. Using Arctos, Berkeley mapper, Google Earth, and Gen-Bank examine Ferruginous Pygmy Owl species Glaucidium brasilianum, specimens MVZ Birds 166038 and MVZ Egg/Nest 8735.

2. And Big Horned sheep species Ovis canadensis nelsoni, Specimens MVZ Mammals 188068, 33619, and 4751.

3. Answer the following questions: a.What will limit these two species if the border wall is built across their range?

b. What will be the result of isolation of different populations? Is it likely that divergence and ultimately speciation will occur?

c. What type of speciation would it be if it occurred? Would it be the same type of speciation for both species?

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Further Investigation

Using the concepts and tools provided in this guide, conduct your own research about the effects of geographic barriers and divergence. Here are a few papers to get you started.

Snapping shrimp speciation:Mathews, L. M. (2006). Cryptic Biodiversity and Phylogeo-graphical Patterns in a Snapping Shrimp Species Complex. Molecular Ecology, 15, 4049-4063.

Jaguars reentering southern states (NM,AZ) :Brown D. E., López González C. A. (2000). Notes on the Occurrences of Jaguars in Arizona and New Mexico. The Southwestern Naturalist, 45, 537-542.

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