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Encyclopedia of Evolution.pdf - Online Reading Center

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disjunct species<br />

have happened? Something, or someone, might have carried<br />

the seeds from Delmarva to the other locations. Neither wind<br />

nor watershed patterns seem to suggest how this could have<br />

occurred. Native Americans use alder bark as a medicine and<br />

may have carried seeds or cuttings with them. The Georgia<br />

population is within the former Cherokee Nation territory,<br />

and the Cherokees are known to have moved south from<br />

the Iroquoian lands <strong>of</strong> what is now the northeastern United<br />

States. Subsequently the Cherokees were relocated by the U.S.<br />

government to Oklahoma in 1838. Could Cherokees have carried<br />

the plants with them, from the Northeast to Georgia to<br />

Oklahoma, connecting all three <strong>of</strong> the existing populations?<br />

This explanation is unlikely to be true, because the three populations<br />

(subspecies) are different enough that they must have<br />

been separated for a long time, much longer than the Cherokee<br />

migrations <strong>of</strong> ca. 1000 and 1838 c.e.<br />

In the case <strong>of</strong> the seaside alder, and in many other cases<br />

as well, the disjunct populations are threatened with extinction.<br />

Not only do small populations have a greater risk <strong>of</strong><br />

extinction, but disjunct populations are frequently not in the<br />

habitats most suitable for them. Oklahoma seaside alders have<br />

been planted, and are surviving well, in Iowa, while they are<br />

barely surviving in Oklahoma, where they can withstand the<br />

summer heat only if the soil is continually wet. This is because<br />

the climatic changes that isolated the disjunct populations in<br />

the first place are now threatening them with extinction. The<br />

Georgia and Delmarva populations are similarly threatened.<br />

In other cases, a species may have a large, central population<br />

and several disjunct populations at some distance away<br />

from the central population. The least tern, for example, is<br />

a shorebird that is abundant in many areas, but the interior<br />

subspecies <strong>of</strong> the least tern (which migrated up the Mississippi<br />

and Arkansas Rivers into Oklahoma) is rare. The white<br />

spruce (Picea glauca) is abundant in the subalpine forests <strong>of</strong><br />

the mountains <strong>of</strong> central North America and the boreal forests<br />

<strong>of</strong> Canada. The limber pine (Pinus flexilis) is fairly common<br />

in the Rocky Mountains. In the Black Hills <strong>of</strong> South Dakota,<br />

relictual populations <strong>of</strong> both the white spruce and the limber<br />

pine survive in conditions that are not cool enough for their<br />

optimal growth. They became stranded in the Black Hills after<br />

the most recent Ice Age and have declined in abundance since<br />

then. The spruce remains abundant in the Black Hills because<br />

it grows well in cool, moist, shaded areas <strong>of</strong> the northern hills.<br />

In contrast, the limber pine has nearly become extinct in the<br />

Black Hills, because it requires cool, dry, sunny conditions,<br />

which are now rare in the Black Hills. The few remaining limber<br />

pines are being crowded and shaded out by the extremely<br />

abundant ponderosa pines (see figure on page 126).<br />

Disjunct populations are frequently the failed dead ends<br />

<strong>of</strong> evolution. They can also be wellsprings <strong>of</strong> evolutionary<br />

novelty. Because they are disjunct, these populations are<br />

reproductively isolated from the larger population, which<br />

allows them to evolve in a different direction. If they have<br />

dispersed from a large, central population, they may also by<br />

chance be genetically different from the original population<br />

(see founder effect). <strong>Evolution</strong> can occur more rapidly in<br />

small populations (see natural selection). If the disjunct<br />

L<br />

P<br />

This is one <strong>of</strong> the last limber pines in the Black Hills <strong>of</strong> South Dakota.<br />

Pinus flexilis (labeled L) is a widespread pine species <strong>of</strong> high mountain<br />

areas in North America, but this population in South Dakota, a remnant<br />

from the most recent Ice Age, is disjunct, small, and heading toward<br />

extinction. Behind the pine is a spruce, Picea glauca (labeled S),<br />

which is also a remnant <strong>of</strong> the most recent Ice Age, and which is<br />

part <strong>of</strong> a relatively abundant population. A ponderosa pine is labeled<br />

P. (Photograph by Stanley A. Rice)<br />

population evolves rapidly, it can produce a new species<br />

which will appear on the scene in what amounts to a geological<br />

instant. Small peripheral (sometimes disjunct) populations<br />

may be largely responsible for the speciation events in the<br />

punctuated equilibria model.<br />

Further <strong>Reading</strong><br />

Schrader, James A., and William R. Graves. “Infraspecific systematics<br />

<strong>of</strong> Alnus maritima (Betulaceae) from three widely disjunct provenances.”<br />

Castanea 67 (2002): 380–401.<br />

———, Stanley A. Rice, and J. Phil Gibson. “Differences in shade tolerance<br />

help explain varying success in two sympatric Alnus species.”<br />

International Journal <strong>of</strong> Plant Sciences, 167 (2006) 979– 989.<br />

S

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