Link: http://insci14.ucsd.edu/~bi178s/fox/foxhistory.

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Hypothesized Evolutionary History and Genetic Characterization
During the mid- to late Pleistocene, the three northern Channel Islands (San
Miguel, Santa Cruz, and Santa Rosa) comprised a single landmass which was
separated from the mainland by a 4-10 km wide channel. It is during this time
(about 16,500 years ago) that gray foxes from the mainland are thought to
have colonized the island by chance dispersal, probably by rafting on floating
debris (Collins 1982). Unique selective pressures in their new habitat probably
caused a reduction in body size and the evolution of genetic differences. These
differences are characteristic of the Channel Island foxes which exists today.
Map reprinted from Wayne et al. 1991
Approximately 11,500 years ago, rising sea levels separated Santa Cruz Island
from the original landmass. As sea levels continued to rise, the remaining island
was further divided into Santa Rosa and San Miguel Islands about 9,500 years
ago (Wayne et al. 1990). Due to this fragmentation, the original fox population
became isolated into three distinct populations, each evolving independently
of one another. The fossil record indicates that foxes first arrived on the southern
islands, on San Clemente Island, approximately 3,400 years ago . Following this
colonization, foxes appeared on San Nicolas Island 2,200 years ago, and on
Santa Catalina Island over 800 years ago. Given the relatively large distance
between northern and southern islands, current speculation is that the foxes
were transported to the southern islands by Native Americans, perhaps as a
source of pelts, as pets, or for religious ceremonies (Collins 1982).

Table 2. Genetic Characterization of Channel Island Foxes^

The genetic analysis done on the six island fox populations has provided the
following evidence for the currently proposed Channel Islands colonization
hypothesis:
-The Santa Cruz Island foxes were most similar to the foxes on Santa Rosa island,
according to the APD value which is based on hypervariable minisatellite DNA.
The APD between these two populations was 52.2% (Gilbert et al. 1990).
-Using hypervariable minisatellite DNA, the foxes on Santa Rosa were shown to

be most closely related to the San Miguel Island foxes. These two populations
had an APD value of 43.8% between them (Gilbert et al. 1990).
-Using hypervariable minisatellite DNA, it was shown that the San Nicolas foxes
were most closely related to the foxes on Santa Catalina. The APD was 55.6%
between these two populations (Gilbert et al. 1990).
-Nei's genetic distance is smallest between the fox populations on the islands of
Santa Rosa and Santa Cruz and the mainland gray fox population (Wayne et
al. 1991). This evidence supports the current colonization hypothesis which
proposes that the San Miguel, Santa Rosa, and Santa Cruz landmass was
originally colonized by mainland gray foxes.
-The level of heterozygosity in the Santa Rosa and Santa Cruz fox populations,
as compared to the heterozygosity of the mainland gray fox, is much higher
than would be predicted by Ht=Ho(1-1/2Ne)t. This equation would predict a
heterozygosity equal to 5.6% of the mainland's fox heterozygosity, but these two
populations are at 42% to 57% of the mainland's fox heterozygosity. This
unexpected result could be due to an increased immigration rate from the
mainland to these islands or possible balancing selection could be favoring the
heterozygotes in these populations (Wayne et al. 1991).
-The higher than expected variability found in the San Miguel fox population
may be due to higher rates of migration between Santa Rosa Island and San
Miguel Island (Wayne et al. 1991). This idea is possible because the water
separating these two islands has been historically lower than it is at the present
time. Higher rates of migration between these two islands could also explain the
observed similarity of hypervariable minisatellite DNA between these two
populations.
-The San Miguel, San Clemente, San Nicolas, and Santa Catalina Islands are
fixed or nearly fixed at the same allozyme loci (Wayne et al. 1991). Such a low
genetic distance suggests a close historical relationship between the foxes of
these islands.
-On San Nicolas Island there has been severe loss of genetic variation due to
bottlenecks and genetic drift. Presumably, the isolation and small population
size on this island has influenced the lack of genetic variation found there
today.
-The lack of allozyme heterozygosity on Santa Catalina Island is very unusual for
a large population size. Furthermore, the Santa Catalina Island fox population
shows a high amount of genetic diversity within its hypervariable minisatellite
DNA and microsatellite DNA. These facts can be sorted out by understanding
that the Santa Catalina fox population experienced severe founder effect
since they were presumably colonized by the San Nicolas foxes which had
already undergone severe drift and possible bottlenecks. This would account
for the lack of allozyme heterozygosity on Santa Catalina. The reason why
Santa Catalina's fox population shows high genetic diversity at the other two
parameters is probably due to the difference in mutation rates. Allozyme loci
have a mutation rate of 10-7 mutations per year while hypervariable

minisatellite loci and microsatellite loci mutate 10-3 and 10-4 times respectively
per year (Wayne et al. 1991 and Wayne 1996). Thus, loci with faster mutation
rates are good indicators of genetic variation in recently diverged populations
(Santa Catalina Island fox population) while slower mutating loci (allozyme
DNA) will show the historic effects of Ne on genetic variability.
-Selection may also have been an important micro evolutionary force in the
maintenance of allozyme variation in the three northern island fox populations
but has been obscured by drift and the founder effect in the three southern
island fox populations (Wayne et al. 1991).