GENETICVARIATION
IN BLACKBEAR POPULATIONSFROMLOUISIANAAND
THE POTENTIALINFLUENCEOF REINTRODUCTIONS
ARKANSAS:EXAMINING
FROMMINNESOTA
A. MILLER,1
DANIEL
Departmentof Fisheriesand WildlifeSciences, VirginiaPolytechnicInstituteand State University,
Blacksburg,VA24061, USA
ERICM. HALLERMAN,
Departmentof Fisheriesand WildlifeSciences, VirginiaPolytechnicInstituteand State University,
Blacksburg,VA24061, USA, email:[email protected]
R. VAUGHAN,
MICHAEL
VirginiaCooperativeFish and WildlifeResearch Unit,Departmentof Fisheriesand WildlifeSciences,
VirginiaPolytechnicInstituteand State University,Blacksburg,VA24061, USA, email:[email protected]
JOHNW. KASBOHM,2
Departmentof Fisheriesand WildlifeSciences, VirginiaPolytechnicInstituteand State University,
Blacksburg,VA24061, USA
Abstract: Using multilocus DNA fingerprinting,we assessed potential genetic effects that may have been caused by translocationof American
black bears (Ursus americanusamericanus)from Minnesotato Louisianaand Arkansas. The bear populationin northeasternMinnesotaexhibits
less within-populationgenetic similarity(similaritywithin = 0.57) than bear populationsin Louisianaand Arkansas(similaritieswithin = 0.74)(P
< 0.001). Populationsin Louisianaand Arkansasare more closely relatedto each other (similaritybetween = 0.53) thanthey are to the population
in Minnesota(similaritybetween = 0.34) (P < 0.001, Mann-Whitneytest). Analysis of band-sharingdata indicatedthat any genetic impactsthat
may have been caused by the translocationswere not statisticallysignificant.
Ursus 10:335-341
Key words: Americanblack bear, Arkansas,DNA fingerprinting,Louisiana,Minnesota,populationgenetics, translocations,Ursus americanus.
The originalrangeof the Americanblackbearincluded
most of North America, including all of Louisiana and
Arkansas.Across this range,the species was dividedinto
16 subspecies (Hall 1981). The Louisianablack bear, U.
a. luteolus, was originallydescribedby Merriam(1893)
on the basis of skull morphology.It historicallyoccupied
easternTexas, all of Louisiana,southernMississippi,and
the southernportion of Arkansas.The American black
bearoccupiedeasternandcentralNorthAmerica,including centraland northernArkansas(Hall 1981).
Populationsof both subspecies declined significantly
in Louisiana and Arkansas during the 1800s and early
1900s due to unregulatedharvestand substantialloss of
forest habitat(Smith and Clark 1994). The bearhunting
season was closed in Arkansasin 1927 due to concerns
aboutlow populationlevels (Smith and Clark 1994). In
1958, because bear populationsremained low, the Arkansas Game and Fish Commission began a restocking
programto increasepopulationsto a level thatwould allow reopeninghuntingseasons (Smith and Clark 1994).
In 1958, 40 bears (U. a. americanus)were capturedin
northeasternMinnesota(LakeandSt. Louis counties)and
releasedin Arkansas. Additionalbears were capturedin
northeasternMinnesotaand released in Arkansasduring
the summersof 1962-68. Release sites for approximately
1Presentaddress: FloridaGame and Fresh WaterFish
Commission, 1239 S.W. 10th Street, Ocala, FL 34474, USA.
2Presentaddress: U.S. Fish and Wildlife Service, Okefenokee
National Wildlife Refuge, Folkston, GA 31537, USA, email:
[email protected]
254 bearsincludedthe OzarkNationalForest(PineyCreek
and White Rock Wildlife ManagementAreas) and the
OuachitaNational Forest (Muddy Creek Wildlife ManagementArea). The precise numberof bearsreleasedat
each site cannotbe determineddue to incompletedocumentation.
Similarly, the Louisiana Departmentof Wildlife and
Fisheries recognized that black bear numberswere low
withintheirstateandbegana restockingprogramin 1964.
Adult bears and cubs were also trappedin northeastern
Minnesota(Cook County),transportedto Louisiana,and
releasedin 2 areas. During 1964-67, approximately130
bears were releasedin the Upper AtchafalayaRiver Basin. Approximately35 bearswere releasedin the Tensas
RiverBasin during1965-66 (Taylor 1971 cited in Pelton
1989).
The Arkansasbearpopulationsincreasedconsiderably
after restocking,to an estimated2,500 bears in western
and northernArkansas,easternOklahoma,and southern
Missouri(SmithandClark1994). Withincreasingpopulation levels, the ArkansasGame and Fish Commission
met its restockingobjectiveandreopenedblackbearhunting in westernandnorthernArkansasin 1980 (Smithand
Clark1994). Huntingwas still closed for a remnantpopulationof nativeblackbearsnearthe WhiteRiverNational
336
Ursus 10:1998
Wildlife Refuge in southeasternArkansas. The current
White River populationis estimatedat 170 bears,but the
populationmay have reacheda low of approximately25
bears during the 1940s (Dellinger 1942 cited in Smith
and Clark 1994).
Recovery recordsof released bears could not provide
conclusive evaluation of the success of the restocking
programsin Louisiana,and populationsremainat notably low levels, as has the amountof suitablebearhabitat.
In 1981, the bear populationswere estimated at 30-50
individualsin the Tensas River area, 30-50 individuals
in the lower AtchafalayaBasin, with an unknownnumber of individualsin the upperAtchafalayaBasin (State
Survey,1981;LouisianaDepartmentof WildlifeandFisheries, cited in Pelton 1989).
Underthe EndangeredSpecies Act of 1973, Congress
definedspecies to include"anysubspeciesof fish or wildlife or plants,andany distinctpopulationsegmentof any
species or vertebratefish or wildlife which interbreeds
when mature"(emphasisadded). Hence, any subspecies
or distinct populationsegment of U. americanusfound
to warrantprotectionbecause of low numberscould be
protected;protectiondid not have to be justified at the
species level. With an increasingconcern over loss of
black bearhabitatand notablylow bear populations,the
U.S. Fish andWildlife Service (USFWS)listed the Louisiana black bear as "threatened"in Januaryof 1992 (57
FederalRegister 588).
During the listing process, questions were raised
about the potential genetic impacts of the bears introduced into Louisiana from Minnesota duringthe 1960s
(Pelton 1991, Smith and Clark 1994). This introduction of U. a. americanus bears into Louisiana could
have altered the putatively unique gene pool of U. a.
luteolus. In making the listing decision, the USFWS
consulted the report by Pelton (1989) that assembled
preliminary genetic and morphological analyses of
black bears from Louisiana, Tennessee, West Virginia,
Virginia, Arkansas, and Minnesota. Genetic marker
frequency data, including protein electrophoresis and
mitochondrial DNA (mtDNA) data, failed to reveal
significant inter-population differentiation, excepting
data from 2 isozyme loci suggesting that Minnesota
populations may be distinct from other populations
(Johns et al. 1989, Zimmerman 1989). However, the
sample sizes for studies reportedin Pelton (1989) were
small (approximately 3 bears/population),limiting the
strength of conclusions. Any genetic impacts of the
introduced bears on the U. a. luteolus genome were
unknown at the time of listing. The objective of this
study, therefore, was to assess any genetic effects of
translocatedbears from Minnesotaon bear populations
in Arkansasand Louisiana.
We thankthe U.S. Fish and Wildlife Service and the
NationalCouncilof the PaperIndustryfor Air andStream
Improvement,Inc. for funding this research. We also
thank the following individuals and agencies for assistancein collectingblood andtissue samples:D. Garshelis,
MinnesotaDepartmentof ForestryandWildlife;D. Goad,
ArkansasGameandFish Commission;M. HurdleandN.
Hunter,White River National Wildlife Refuge (NWR);
H. Jacobsonand T. White, Mississippi State University;
T. Edwards,Tensas River NWR; M. Pelton and personnel, University of Tennessee; R. Pace, Louisiana State
University; and K. Shropshire,Mississippi Department
of Wildlife, Fisheries, and Parks. S. Fain, USFWS National Fish and Wildlife Forensics Laboratory,provided
access to unpublishedlaboratorymethods and data. In
addition, we thank B. Lubinski and J. Allen, Virginia
Polytechnic Instituteand State University, (VPISU) for
laboratoryassistance, S. Miller, VPISU, and T. Eason,
University of Tennessee, for field assistance, and J.
Kasbohm,VPISU, for his guidance and assistance collecting samples. We also thankan anonymousreviewer
for helpful comments.
METHODS
SampleCollection
We collected blood and tissue samples from 7 black
bear populationsin Arkansas,Louisiana,andMinnesota
(Fig. 1). Bears were trappedand immobilized for blood
collection using standard procedures (Kasbohm and
Vaughan1993); additionaltissue sampleswere collected
from hunter-killedand road-killedbears. Blood and tissue samples were stored at -60?C or mixed with a sodium dodecyl sulphate buffer for storage at ambient
temperatures(Kirby 1992).
DNAFingerprinting
We extracted whole genomic DNA from blood and
tissue using standardproteinaseK digestion and phenol/
chloroformextractiontechniques(Sambrooket al. 1989).
PurifiedDNA was digested using the restrictionenzyme
HinfI, andthe DNA sampleswere subjectedto TBE agarose gel electrophoresis according to Sambrook et al.
(1989). We used lambda DNA cut with BstElI (New
EnglandBiolabs, Beverly, Mass.),placedin severallanes
across each gel, as a molecular weight marker. DNA
fragments were transferredfrom the agarose gel to a
Magnagraphnylon membrane(MSI, Westboro, Mass.)
BLACKBEARGENETICS* Miller et al.
via Southernblotting (Southern 1975). We hybridized
Southernblots using Jeffreys' minisatelliteprobe 33.15
(Jeffreys et al. 1985) tagged with chemiluminescent
agents (CellmarkDiagnostics, Germantown,Md.). After hybridizationusing the specific protocol established
by Cellmark Diagnostics, we exposed each membrane
to Kodak XAR5 X-omat X-ray film (Rochester,N.Y.).
The hybridizedregionson each membranewere observed
as bandingpatternson the autoradiograph.
and Analysisof
Interpretation
DNAFingerprints
We quantifiedthe degree of relatednessbetween any 2
individualbearsin termsof the numberof bandsof equal
electrophoreticmobility exhibitedby both individualsin
the 2.3-7.2 kilobaserangeof molecularweights. A measure of phenotypic similarity, S, was calculated for all
pairingsof individuals,allowing for comparisonswithin
and between populationsexaminedon the same autoradiograph(Brufordet al. 1992). We calculatedS as:
S = 2nyl(n+ny), where
ny = the numberof bands exhibitedby both
individuals,
nx= the total numberof bands exhibitedby
individualx, and
ny = the total numberof bandsexhibitedby
individualy.
Median similarityvalues of S were comparedwithin
and between Arkansas,Louisiana,and Minnesotapopulations. Because datawere non-normallydistributed,we
used medianinsteadof mean values and non-parametric
statistical procedures. Mann-Whitney tests were performed to determinesignificantdifferences.
RESULTS
We screened genetic variability among DNA
samples of 103 bears from 7 populations using DNA
fingerprintingtechniques (Table 1). Median estimated
similarity values within populations (reported along
the diagonal of Table 2) ranged from 0.57 (Cook
County, Minn.) to 0.81 (White River NWR). Median
estimated similarity values between populations
ranged from 0.19 (White River NWR versus Ouachita
National Forest) to 0.70 (White River NWR versus
Tensas NWR) (Table 2, Fig. 1).
Within-populationsimilarityvalues (0.57 versus 0.74)
within the Minnesota and Arkansas-Louisianapopulations were different (P < 0.001, Mann-Whitney test).
Among the 3 Arkansas populations, estimated genetic
similarity was higher (P < 0.001, Mann-Whitney test)
337
within White River NWR (0.81) than within the Ozark
and OuachitaNational Forest populations(0.63 pooled)
(Table2). Among the 3 Louisianapopulations,estimated
genetic similaritywas lower (P < 0.001, Mann-Whitney
test) within the Upper Atchafalaya Basin (0.67) than
within the Tensas River NWR and Lower Atchafalaya
Basin populations(0.78 pooled).
The estimated median genetic similaritybetween the
Cook County, Minnesota population and each population fromLouisianaandArkansaswas 0.34 (5 inter-population comparisons). The estimated median genetic
similaritybetween the Louisiana and Arkansaspopulations was 0.53 (6 inter-populationcomparisons)(Table
3). These 2 between-populationsimilarityvalues were
different(P < 0.001, Mann-Whitneytest).
DISCUSSION
GeneticVariation
withinPopulations
Band-sharingdataindicatedthatthe bearpopulationin
Cook County, Minnesotaexhibited more genetic diversity thanbearpopulationsfrom Louisianaand Arkansas.
The Minnesota populationis much larger than the isolated Louisianaor Arkansaspopulations(Jonkel 1978),
and largerpopulationstend to show more genetic variation than smaller, isolated populations (Allendorf and
Leary 1986, Hartl 1988).
Among the 3 populationsin Arkansas,the WhiteRiver
NWR populationexhibitedthe highestwithin-population
genetic similarity. Even though the White River NWR
populationmay have declined to approximately25 individuals in the 1940s (Dellinger 1942 cited in Smith and
Clark 1994), this populationwas not partof the restocking programin Arkansas. The populationremains at a
low numberrelativeto the Ozarkand OuachitaNF populations(SmithandClark1994), andtheWhiteRiverNWR
populationmay have lost genetic variationdue to a genetic bottleneck and associated random genetic drift
(Allendorf 1986, Hartl 1988). Smith and Clark (1994)
estimatethatapproximately2,500 bearsnow inhabitwestern and northernArkansasand portionsof easternOklahomaandsouthernMissouri.Restockingeffortsmayhave
reducedany effects of randomgenetic driftandbe partly
responsible for the higher genetic diversity within the
Ozarkand OuachitaNF populations. The rapidrecovery
fromany genetic bottleneckmay have minimizedassociated losses of genetic diversity (Nei et al. 1975).
Among the 3 black bearpopulationsin Louisiana,the
Tensas River NWR and Lower AtchafalayaBasin populations showed less genetic variation.This may be due to
338
Ursus 10:1998
Table 1. DNA fingerprint banding sharing patterns within black bear populations in Minnesota, Arkansas, and Louisiana (Hinfl
restriction digests, Jeffreys' 33.15 probe). Due to the non-normality of the data, values are given as medians, with ranges
given in parentheses. Samplesizes (n) given referto the numberof bears sampled/population.A shared band is one
exhibited by all individuals sampled within a given population; a polymorphic band is one not exhibited by all individuals in
a given population.
(n)
Populationa
Bands/individual
(range)
Sharedbands
(range)
COOK (31)
OZAR (8)
OUAC (8)
WRIV (17)
TENS (16)
UATC (12)
LATC (11)
22.0
27.0
27.0
26.0
19.5
25.0
14.5
3.0
9.0
10.0
12.0
9.0
5.0
7.0
(14-33)
(23-33)
(21-32)
(20-38)
(10-26)
(15-33)
(12-27)
bands
Polymorphic
(range)
46.5
39.0
35.5
25.0
15.5
37.0
23.0
(1-6)
(6-12)
(9-11)
(11-23)
(5-13)
(2-8)
(5-9)
(28-55)
(31-47)
(28-43)
(19-27)
(13-37)
(27-40)
(18-28)
Totalbands
(range)
49.5
48.0
45.5
38.0
27.5
42.5
30.0
(32-56)
(43-53)
(39-52)
(31-48)
(20-42)
(29-47)
(23-37)
a Locales of genetic sampling(n = sample size): COOK = Cook County, Minn.; OZAR = OzarkNationalForest, Ark.; OUAC = Ouachita
National Forest, Ark.; WRIV = White River National Wildlife Refuge, Ark.; TENS = Tensas River National Wildlife Refuge, La.; UATC =
Upper AtchafalayaBasin, La.; LATC = Lower AtchafalayaBasin, La.
a genetic bottleneckwith associatedrandomgenetic drift
(Hartl1988), given thatthese populationswere each estimated to have only 30-50 bears in 1981 (State Survey,
1981, Louisiana Departmentof Wildlife and Fisheries
cited in Pelton 1989). Even though approximately35
bears were translocatedto the Tensas River NWR populationduringthe restockingprogram,Nowak (1986 cited
in Pelton 1989) speculatedthatnone of the releasedbears
survivedor remainedin the area. The LowerAtchafalaya
Basin populationwas not partof the restockingprogram.
The UpperAtchafalayaBasin exhibitedthe most withinpopulationgenetic variation;restockingeffortsmay have
reducedthe impactsof randomgenetic drift(Hartl1988)
andbe partlyresponsiblefor the highergenetic diversity.
The possibility that translocatedblack bears may have
affected the U. a. luteolus gene pool, therefore,cannot
be excluded.
GeneticVariationbetweenPopulations
Bear populationsin Louisianaand Arkansasare more
closely relatedto each otherthanthey are to the population in Minnesota.Estimatedgenetic similaritiesbetween
black bear populationsin Louisianaand Arkansaswere
higher than the estimated genetic similarities between
the Cook County,Minnesota,populationandeach population in Louisiana and Arkansas (P < 0.001, MannWhitney test). Although restocking efforts may have
influenced levels of genetic variability within certain
Louisiana and Arkansas bear populations, any genetic
impacts caused by these translocationswere not significantenoughto alterthe overallgenetic similaritybetween
populations.
Within Arkansas, the White River NWR population
was more similarto the OzarkNF populationthanto the
OuachitaNF population. Bears use narrowriver corri-
valuesforgeneticsimilaritiesof blackbearpopulationsin Minnesota,Arkansas,and
Table2. Matrixof medianband-sharing
Louisiana(Hinflrestrictiondigests,Jeffreys'33.15probe).Valuesalongthe diagonalrepresentestimatedgeneticsimilarities
withinpopulations;values below the diagonalrepresentestimatedgenetic similaritiesbetweenpopulations. Asterisks
indicateabsence of similarityestimatesfor pairsof populationsforwhichDNAfingerprintswere not observedon the same
Southernblot.
Populationa
COOK
OZAR
OUAC
WRIV
TENS
UATC
LATC
COOK
OZAR
OUAC
WRIV
TENS
UATC
LATC
0.57
0.30
0.23
0.39
0.41
0.29
****
0.63
0.43
0.405
****
****
****
0.635
0.19
****
****
****
0.81
0.70
****
****
0.78
0.56
****
0.67
0.50
0.78
0.78
a COOK = Cook County, Minn.; OZAR = OzarkNational Forest, Ark.; OUAC = OuachitaNational Forest, Ark.; WRIV = White River
National Wildlife Refuge, Ark.; TENS = Tensas River National Wildlife Refuge, La.; UATC = Upper AtchafalayaBasin, La.; LATC =
Lower AtchafalayaBasin, La.
* Miller et al.
BLACKBEARGENETICS
339
21 U. UpperAtchafalayaBasin, LA
21L LowerAtchafalayaBasin, LA
22. Tensas RiverNWR,LA
23. WhiteRiverNWR,AR
24. OuachitaNF,AR
25. OzarkNF,AR
27. Cook County,MN
Fig. 1. Median band-sharing values for genetic similarities within and between black bear populations in Minnesota,
Arkansas, and Louisiana (Hinfl restriction digests, Jeffrey's 33.15 probe). Numbers enclosed in circles are arbitraryand
represent populations, numbers enclosed in triangles represent genetic similarities within populations, and numbers
enclosed in rectangles represent genetic similarities between populations.
dors for dispersalor migration(Pelton 1989) and potentially can travel between the OzarkNF and White River
NWR (approximately250 km) using the White River
corridor,providinga means for gene flow between these
2 populations. In contrast,travel between the Ouachita
NF andWhite River NWR (approximately175 km) may
be limited by migrationbarriers,possibly including the
ArkansasRiver, the city of Little Rock, and several major interstates.
Estimatedgenetic similaritybetween the White River
NWR, Arkansas, and Tensas River NWR, Louisiana,
populations was high (0.70), suggesting gene flow be-
tween these 2 populations (Pelton 1989). Given the
small sizes of the White River NWR and Tensas River
NWR populations, sufficient migration would be necessary to keep the 2 populationsfrom diverging genetically due to randomgenetic drift;the actualdivergence
of the populationswould be a functionof the numberof
migrantsper generationand of means and variancesof
allele frequencies (Allendorf and Phelps 1981).
StatisticalConsiderations
Populationdatacollected with DNA fingerprintingdo
not providea completecharacterizationof genetic varia-
340
Ursus 10:1998
Table3. Medianband-sharing
valuesforgeneticsimilarities
of black bear populationsin Minnesota,Arkansas,and
Louisiana
Population
Median
band-sharing
Within individualpopulations:(P < 0.001)
Minnesota (n = 31)
Louisiana-Arkansas(n = 72)
0.57
0.74
Between individualpopulations: (P < 0.001)
Minnesotavs. Louisiana-Arkansas(n = 77)
Louisiana-Arkansas(n = 56)
0.34
0.53
tion in termsof allele frequencydistributionsbecauseneitherthe numberof loci nor the locus affiliationof alleles
is directlyobservable(Scott andWilliams 1994). Use of
certain traditionalpopulationgenetic statistics was not
found defensiblein the context of small bearpopulations
subjectto restocking.
Inferences regardingaverage heterozygosity and genetic distancebased on DNA fingerprintingdatarequire
2 assumptionsbeyondthoserequiredfor single-locusdata
(Jin and Chakraborty1994). First,the Hardy-Weinberg
rule is assumedfor each population,but the assumption
thatmigrationis unimportantclearlyis violatedin a small
populationsubjectto reintroduction.Imprecisionin estimatingbandfrequenciesdue to samplingerroralso compromises tests based on Hardy-Weinbergassumptions.
Second,co-migrationof bandsfromdistinctminisatellite
loci and the assumptionthat they representthe same allele has an unknowneffect on estimationof population
genetic statistics.
Band-sharingdata often are analyzed in the genetics
literatureusing classicalstatisticalproceduresthatassume
normaldata (Gilbertet al. 1990, Westneat 1990, Wayne
et al. 1991, Triggs et al. 1992). However, non-parametric proceduresshould be used when data do not come
from a normaldistribution(Hollanderand Wolfe 1973).
Median values provide a more reliablerepresentationof
non-normaldistributionsthanmeanvalues;therefore,we
used median values and non-parametrictests for differences in our analysis
Given the pair-wisenatureof the similarityindex calculation, band-sharingdata from each DNA fingerprint
represent a collection of non-independentdata points
(Wayneet al. 1991). Underthe Mann-Whitneytest,however,datapointsareassumedto be independent.Although
the independenceassumptionis violated, the currentgenetics and statisticsliteratureoffer no alternativestatistical methods for handlingdependentdata. Although the
robustness of the Mann-Whitney test to non-indepen-
denceis unknown,afterconsultationwith statisticiansand
geneticists, we chose to use the non-parametricproceduresdiscussed for data analysis.
Lynch (1990) provided a method for partitioningthe
comsimilarityindexinto within-andbetween-population
but
are
needed
to
make
ponents,
generous assumptions
the transition. Stephenset al. (1992) presenteda method
for calculatingaverageheterozygosityusing band-sharing data,providedthat allele frequenciescan be inferred
from such data. Hence, these 2 statisticalmethodswere
not used in this study due to questionsregardingthe satisfaction of the underlyingassumptionseach required.
Conclusions
Any genetic impactsthatmay have been causedby the
translocationswere not statisticallysignificant. The bear
Minnesotaexhibitsless withinpopulationof northeastern
population genetic similarity than bear populations in
Louisiana and Arkansas, and populationsin Louisiana
and Arkansasare more closely relatedto each otherthan
they are to the populationin Minnesota.
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