Cavenaugh, Carrie F.

Johnson, Ronald L.

UARK (University of Arkansas )

Journal of the Arkansas Academy of Science Volume 57 Article 32 2003 Genetic Similarity of Shadow and Arkansas Basses (Ambloplites) Determined by Mitochondrial DNA Analysis Ronald L. Johnson Arkansas State University Carrie F. Cavenaugh Arkansas State University Follow this and additional works at: https://scholarworks.uark.edu/jaas  Part of the Zoology Commons Recommended Citation Johnson, Ronald L. and Cavenaugh, Carrie F. (2003) "Genetic Similarity of Shadow and Arkansas Basses (Ambloplites) Determined by Mitochondrial DNA Analysis," Journal of the Arkansas Academy of Science: Vol. 57 , Article 32. Available at: https://scholarworks.uark.edu/jaas/vol57/iss1/32 This article is available for use under the Creative Commons license: Attribution-NoDerivatives 4.0 International (CC BY-ND 4.0). Users are able to read, download, copy, print, distribute, search, link to the full texts of these articles, or use them for any other lawful purpose, without asking prior permission from the publisher or the author. This General Note is brought to you for free and open access by ScholarWorks@UARK. It has been accepted for inclusion in Journal of the Arkansas Academy of Science by an authorized editor of ScholarWorks@UARK. For more information, please contact scholar@uark.edu. 206Genetic Similarity of Shadow and Ozark Basses [Ambloplites) asDetermined by Mitochondrial DNA AnalysisRonald L.Johnson* and Carrie F. CavenaughArkansas State UniversityDepartment of Biological SciencesState University, AR 72467*Corresponding AuthorThe rock basses, Ambloplites, contain four nominalspecies and represent one of eight genera within the sunfishfamily (Centrarchidae). The shadow bass (A. ariommusViosca) occurs in the southern plains states east and west ofthe Mississippi River. The range of the shadow bass west ofthe Mississippi River is limited to Arkansas and parts ofsouthern Missouri and eastern Oklahoma. Several Arkansasriver drainages contain the shadow bass including the Red,Ouachita, Arkansas, Illinois,Little Red, Strawberry, Spring,Black, and St. Francis river drainages (Robison andBuchanan, 1988). The Ozark bass {A. constellatus Cashnerand Suttkus) is found in streams of the White River drainagein northern Arkansas and southern Missouri and in theBuffalo River (Robison and Buchanan, 1988). Thegeographic ranges of the shadow and Ozark basses areproximal to one another. The Roanoke bass (A. cavifronsCope) is limited to Virginia and North Carolina (Cashnerand Jenkins, 1982). Only the rock bass, A. rupestrisRafinesque, has an extensive range throughout much of theMississippi River drainage.Several morphological and anatomical featuresdistinguish the shadow and Ozark basses including blotchypigmentation versus irregular rows of spots, a deeper versusa more slender body depth, and the presence of less than ormore than 41 lateral line scales (Robison and Buchanan,1988). Genetic analysis of Ambloplites is limited to a singleallozyme study. Koppleman et al. (2000) could notgenetically distinguish between the shadow and rock basses,yet identified fixed allelic differences at two of 41 locibetween the Ozark and shadow basses. No direct DNAcomparisons have been performed on this genus. One ofour research goals was to perform a genetic comparison ofthe Ambloplites species native to Arkansas (shadow andOzark basses) using restriction endonuclease digestion ofintact mitochondrial DNA (mtDNA). A second goal was toestimate the time of divergence using a commonly usedmtDNA molecular clock (Brown et al., 1979).Shadow bass were collected from the South Fork of theSpring River (n =22), the Spring River (n =28), and theCaddo River (n = 2), while Ozark bass (n = 24) werecollected from the Buffalo River. Mitochondria and mtDNAwere isolated and analyzed from liver tissue usingtechniques described by Johnson et al. (2002). PurifiedmtDNA was digested for 7 h at 37° C using 14 restrictionendonucleases {BamHl, BgH, BgUl, Csp45l, Dral, EcoRl,EcoRV, MM, Pstl, Pvull, Sail, Seal, Xbal, Xhol), underconditions recommended by the supplier (Promega Corp.).Fragment sizes generated were determined through the useof a least-squares fitprogram (Schaeffer and Sederoff, 1981).Variables measured included genome size, percentagegenome analyzed, nucleon diversity (h; Nei and Tajima,1981) and nucleotide sequence divergence (Nei and Li,1979). Times of divergence were estimated from observedlevels of sequence divergence using a rate of sequencedivergence of 2% per million years (Brown et al., 1979). Aphenogram was constructed by the unweighted pair groupmethod (UPGMA; Sokal and Sneath, 1963) using matricesof distance values with NTSYS-PC: Numerical Taxonomyand Multivariate Analysis System (Rohlf, 1990) as wellas aninferred phylogenetic tree using the Dollo parsimonyalgorithm in Phylogeny Inference Package (PHYLIP;Felsenstein, 1993). The northern largemouth bass(Micropterus salmoides salmoides Lacepede) and the greensunfish (Lepomis cyanellus Rafinesque), basal members ofother centrarchid genera, were used for comparison ofhigher taxonomic relationships (Branson and Moore, 1962;Avise and Smith, 1977).Mean genome size ranged from 16,676 (± 337 SE) basepairs for shadow bass to 17,170 (± 292 SE) base pairs forOzark bass. These estimates are similar to values obtainedfor Lepomis (range 16,639 to 16,958; Johnson and Williams,2003) and slightly lower than that obtained for Micropterus(range of 17,346 to 17,779; Johnson et al., 2002).The total number of restriction fragments generatedusing 13 restriction endonucleases was 43, with 31 fragmentsidentified for the Ozark bass and 32 fragments for theshadow bass. Three of the restriction endonucleases did notidentify recognition sequences within individual species(BamHl, Ozark bass, shadow bass; Sail, shadow bass; andXhol,Ozark bass and shadow bass) [Table 1]. The dominanthaplotype (A) for the shadow bass was found in all threepopulations sampled (Caddo, South Fork and Spring rivers).Two additional haplotypes (B and C) were identified forsingle individuals from the South Fork (B) and the SpringRiver (C). Haplotype diversity for shadow bass of the SouthFork was 0.15 with a diversity of 0.07 for the Spring RiverJournal of the Arkansas Academy of Science, Vol. 57, 2003206Journal of the Arkansas Academy of Science, Vol. 57 [2003], Art. 32Published by Arkansas Academy of Science, 2003207Genetic Similarity of Shadow and Ozark Basses {Ambloplites) as Determined by Mitochondria! DNAAnalysisTable 1. Composite haplotypes ofA. ariommus, A.constellates,L. cyanellus and M. salmoides collected from northeastArkansas. Restriction endonucleases are in columns asfollows: BamHI, BgH, BgUl, Csp45l, Dral, EcoRl, EcoRV,Mlul,Pstl, Pvull, SaH, Seal, Xbal, and Xhol.Species Genotype Assignment nA. ariommus A AAAAAAAAAAAAA 50A.ariommus B AAAAABAAAAAAA 1A.ariommus C AAAAAABAAAAAA 1A.constellatus ABAABABBABBBA 21M. salmoides BCABCACCBCCCB 1L. cyanellus CDACDCDDCBDDC 1population. Haplotype B had a unique restriction site for theendonuclease EcoRV, whereas haplotype C had a uniquerestriction site for Mlul. There was a single haplotype foundwithin the Ozark bass sampled (h =0.00). Unique restrictionprofiles between the Ozark and shadow basses wereidentified for the following restriction endonucleases: BgH,Dral, Pstl, Sail, Seal, and Xbal.Estimated nucleotide sequence divergence, standarderror, and time of divergence for Ambloplites, Lepomis, andMicropterus are found in Table 2. Divergence values rangedfrom 0.0612 for the shadow and Ozark basses to 0.3216 forthe largemouth bass and green sunfish. The divergencevalues for the shadow and Ozark basses are lower than thatbund for congeneric members within the genera Lepomisand Micropterus (Johnson and Williams,2003; Johnson et al.,2002). Data indicated that Ambloplites is more similar to theargemouth bass than to the green sunfish. Estimated timeofdivergence for the two species of Ambloplites studied was3.07 million years ago (Table 4). Unfortunately, there is noreported fossil record of Ambloplites on which to base acomparison. Itmust be noted that the molecular clock of2%)er million years has not been consistent through all taxaGrewe et al., 1990). Additionally, these estimates are basedon few populations within each species and therefore thevalues obtained are at best estimations of the divergence>eriods between these particular populations rather than thespecies as a whole.Construction of a cladogram generated utilizing the)ollo parsimony algorithm program revealed thatAmbloplites was most similar to Micropterus and less so toLepomis (Fig. 1). A UPGMA dendrogram derived from theaverage genetic distances between species was consistentwith that above. A cophenetic correlation of 0.76 wasTable 2. Mitochondrial DNA sequence divergence (abovediagonal) and estimated time of divergence in millions ofyears (below diagonal) for Ambloplites, Lepomis andMicropterus studied. Standard errors of the mean inparentheses beneath the means.Species 12 3 41. A.ariommus***0.0615 0.2044 0.1675***(0.0014) (0.0003) (0.0030)2. A. constellates 3.07***0.2121 0.1371(0.01)***(0.0004) (0.0006)3. L.cyanellus 10.22 10.66***0.3216(0.01) (0.01)***(0.0011)4. M.salmoides 8.37 6.86 16.08***(0.01) (0.01) (0.01) ***obtained, indicative of a suitable fit between the treegenerated and the data matrix (Rohlf, 1990). In contrast,Avise and Smith (1977) hypothesized that Lepomis wasintermediate to Micropterus and Ambloplites based uponallozyme analysis of several centrarchid genera. However,Avise and Smith (1977) sampled the rock bass as comparedto the shadow and Ozark basses, and they used the averagegenetic distances of ten Lepomis species rather than lookingat a specific species (green sunfish). Future study of thegenus Ambloplites at the population, subspecies, and specieslevels is required to further our understanding of thehistorical biogeography of this genus. Additional populationdata may alter the proposed relationships of these taxa.We would like to thank J. Quinn, K. Shirley, M. Wine,C. Hutts, T. Sanders and L. Mommsen for technicalassistance and for assistance in the collection ofspecimens.This work was supported by grants from the SiloUndergraduate Research Foundation and Arkansas StateUniversity.Literature CitedAvise, J. C, and M. H. Smith. 1977. Gene frequencycomparisons between sunfish (Centrarchidae)populations at various stages of evolutionarydivergence. Syst. Zool. 26: 319-335Branson, B. A., and G. A. Moore. 1962. The lateraliscomponents of the acoustico-lateralis system in thesunfish family Centrarchidae. Copeia 1962: 1-108.Journal of the Arkansas Academy of Science, Vol. 57, 2003207Journal of the Arkansas Academy of Science, Vol. 57 [2003], Art. 32https://scholarworks.uark.edu/jaas/vol57/iss1/32208Ronald L.Johnson and Carrie F. CavenaughL. cyanellusM.salmoidesA. constellatusA. ariommus 'CA.ariommus 'B'A.ariommus 'A'Fig. 1. Cladogram of character state matrices for presence/absence ofpresumptive restriction sites utilizing the Dolloparsimonyalgorithm. One parsimonious tree (22 steps) was generated.Brown, W. M.,M.George, Jr., and A. C. Wilson. 1979.Rapid evolution of animal mitochondrial DNA. Proc.Nat. Acad. Sci. 76:1967-1971.Cashner, R. C,and R.E.Jenkins. 1982. Systematics of theRoanoke bass, Ambloplites cavifrons. Copeia 1982:581-594.Felsenstein, J. 1993. Phylip (Phylogeny Inference Package).Version 3.51. University of Washington, Seattle.Grewe, P. M.,N. Billington, and P. D.N. Hebert 1990.Phylogenetic relationships among members ofSalvelinus inferred from mitochondrial DNAdivergence. Can. J. Fish. Aq.Sci. 47:984-991.Johnson, R. L., and K. A. Williams. 2003. Geneticrelationships of some common Arkansas freshwatersunfishes (Centrarchidae: Lepomis) inferred fromrestriction endonuclease analysis of mitochondrialDNA.J. Arkansas Acad. Sci. 57:(in press).Johnson, R. L., J. B. Magee, and T. A. Hodge. 2002.Phylogenetics of freshwater black basses(Centrarchidae: Micropterus) inferred from restrictionendonuclease analysis of mitochondrial DNA.Biochem. Gen. 39:395-496.Koppleman, J. B., C.M.Gale, andj. S. Stanovick. 2000.Allozyme and morphological variation among threenominal species of Ambloplites (Centrarchidae)inhabiting the Ozarks Region. Trans. Am. Fish. Soc.129:1134-1149.Nei, M., and W. H. Li. 1979. Mathematical models forstudying genetic variation in terms of restrictionendonucleases. Proc. Nat. Acad. Sci. 76:5269-5273.Nei, M., and F. Tajima. 1981. DNA polymorphismsdetectable by restriction endonucleases. Genetics97:147-165.Robison, H. W, and T. M. Buchanan. 1988. Fishes ofArkansas. University of Arkansas Press, Fayetteville.563 pp.Rohlf, F. F. 1990. NTSYS-PC: Numerical Taxonomy andMultivariate Analysis System. Version 1.60. ExeterPublishing, Setauket.Schaeffer, H. E., and R. R. Sederoff. 1981. Improvedestimation on DNA fragment lengths from agarose gels.Ann. Biochem. 115:113-122.Sokal, R. R., and P. H. A. Sneath. 1963. Principles ofnumerical taxonomy. Freeman, San Francisco. 359 pp.Journal of the Arkansas Academy of Science, Vol. 57, 2003208Journal of the Arkansas Academy of Science, Vol. 57 [2003], Art. 32Published by Arkansas Academy of Science, 2003

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Genetic Similarity of Shadow and Arkansas Basses (Ambloplites) Determined by Mitochondrial DNA Analysis

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Journal of the Arkansas Academy of ScienceVolume 57 Article 322003Genetic Similarity of Shadow and Arkansas Basses(Ambloplites) Determined by MitochondrialDNA AnalysisRonald L. JohnsonArkansas State UniversityCarrie F. CavenaughArkansas State UniversityFollow this and additional works at: http://scholarworks.uark.edu/jaasPart of the Zoology CommonsThis article is available for use under the Creative Commons license: Attribution-NoDerivatives 4.0 International (CC BY-ND 4.0). Users are able toread, download, copy, print, distribute, search, link to the full texts of these articles, or use them for any other lawful purpose, without asking priorpermission from the publisher or the author.This General Note is brought to you for free and open access by ScholarWorks@UARK. It has been accepted for inclusion in Journal of the ArkansasAcademy of Science by an authorized editor of ScholarWorks@UARK. For more information, please contact scholar@uark.edu.Recommended CitationJohnson, Ronald L. and Cavenaugh, Carrie F. (2003) "Genetic Similarity of Shadow and Arkansas Basses (Ambloplites) Determinedby Mitochondrial DNA Analysis," Journal of the Arkansas Academy of Science: Vol. 57 , Article 32.Available at: http://scholarworks.uark.edu/jaas/vol57/iss1/32206Genetic Similarity of Shadow and Ozark Basses [Ambloplites) asDetermined by Mitochondrial DNA AnalysisRonald L.Johnson* and Carrie F. CavenaughArkansas State UniversityDepartment of Biological SciencesState University, AR 72467* Corresponding AuthorThe rock basses, Ambloplites, contain four nominalspecies and represent one of eight genera within the sunfishfamily (Centrarchidae). The shadow bass (A. ariommusViosca) occurs in the southern plains states east and west ofthe Mississippi River. The range of the shadow bass west ofthe Mississippi River is limited to Arkansas and parts ofsouthern Missouri and eastern Oklahoma. Several Arkansasriver drainages contain the shadow bass including the Red,Ouachita, Arkansas, Illinois,Little Red, Strawberry, Spring,Black, and St. Francis river drainages (Robison andBuchanan, 1988). The Ozark bass {A. constellatus Cashnerand Suttkus) is found in streams of the White River drainagein northern Arkansas and southern Missouri and in theBuffalo River (Robison and Buchanan, 1988). Thegeographic ranges of the shadow and Ozark basses areproximal to one another. The Roanoke bass (A. cavifronsCope) is limited to Virginia and North Carolina (Cashnerand Jenkins, 1982). Only the rock bass, A. rupestrisRafinesque, has an extensive range throughout much of theMississippi River drainage.Several morphological and anatomical featuresdistinguish the shadow and Ozark basses including blotchypigmentation versus irregular rows of spots, a deeper versusa more slender body depth, and the presence of less than ormore than 41 lateral line scales (Robison and Buchanan,1988). Genetic analysis of Ambloplites is limited to a singleallozyme study. Koppleman et al. (2000) could notgenetically distinguish between the shadow and rock basses,yet identified fixed allelic differences at two of 41 locibetween the Ozark and shadow basses. No direct DNAcomparisons have been performed on this genus. One ofour research goals was to perform a genetic comparison ofthe Ambloplites species native to Arkansas (shadow andOzark basses) using restriction endonuclease digestion ofintact mitochondrial DNA (mtDNA). A second goal was toestimate the time of divergence using a commonly usedmtDNA molecular clock (Brown et al., 1979).Shadow bass were collected from the South Fork of theSpring River (n =22), the Spring River (n =28), and theCaddo River (n = 2), while Ozark bass (n = 24) werecollected from the Buffalo River. Mitochondria and mtDNAwere isolated and analyzed from liver tissue usingtechniques described by Johnson et al. (2002). PurifiedmtDNA was digested for 7 h at 37° C using 14 restrictionendonucleases {BamHl, BgH, BgUl, Csp45l, Dral, EcoRl,EcoRV, MM, Pstl, Pvull, Sail, Seal, Xbal, Xhol), underconditions recommended by the supplier (Promega Corp.).Fragment sizes generated were determined through the useof a least-squares fitprogram (Schaeffer and Sederoff, 1981).Variables measured included genome size, percentagegenome analyzed, nucleon diversity (h; Nei and Tajima,1981) and nucleotide sequence divergence (Nei and Li,1979). Times of divergence were estimated from observedlevels of sequence divergence using a rate of sequencedivergence of 2% per million years (Brown et al., 1979). Aphenogram was constructed by the unweighted pair groupmethod (UPGMA; Sokal and Sneath, 1963) using matricesof distance values with NTSYS-PC: Numerical Taxonomyand Multivariate Analysis System (Rohlf, 1990) as wellas aninferred phylogenetic tree using the Dollo parsimonyalgorithm in Phylogeny Inference Package (PHYLIP;Felsenstein, 1993). The northern largemouth bass(Micropterus salmoides salmoides Lacepede) and the greensunfish (Lepomis cyanellus Rafinesque), basal members ofother centrarchid genera, were used for comparison ofhigher taxonomic relationships (Branson and Moore, 1962;Avise and Smith, 1977).Mean genome size ranged from 16,676 (± 337 SE) basepairs for shadow bass to 17,170 (± 292 SE) base pairs forOzark bass. These estimates are similar to values obtainedfor Lepomis (range 16,639 to 16,958; Johnson and Williams,2003) and slightly lower than that obtained for Micropterus(range of 17,346 to 17,779; Johnson et al., 2002).The total number of restriction fragments generatedusing 13 restriction endonucleases was 43, with 31 fragmentsidentified for the Ozark bass and 32 fragments for theshadow bass. Three of the restriction endonucleases did notidentify recognition sequences within individual species(BamHl, Ozark bass, shadow bass; Sail, shadow bass; andXhol,Ozark bass and shadow bass) [Table 1]. The dominanthaplotype (A) for the shadow bass was found in all threepopulations sampled (Caddo, South Fork and Spring rivers).Two additional haplotypes (B and C) were identified forsingle individuals from the South Fork (B) and the SpringRiver (C). Haplotype diversity for shadow bass of the SouthFork was 0.15 with a diversity of 0.07 for the Spring RiverJournal of the Arkansas Academy of Science, Vol. 57, 2003206Journal of the Arkansas Academy of Science, Vol. 57 [2003], Art. 32Published by Arkansas Academy of Science, 2003207Genetic Similarity of Shadow and Ozark Basses {Ambloplites) as Determined by Mitochondria! DNAAnalysisTable 1. Composite haplotypes ofA. ariommus, A.constellates,L. cyanellus and M. salmoides collected from northeastArkansas. Restriction endonucleases are in columns asfollows: BamHI, BgH, BgUl, Csp45l, Dral, EcoRl, EcoRV,Mlul,Pstl, Pvull, SaH, Seal, Xbal, and Xhol.Species Genotype Assignment nA. ariommus A AAAAAAAAAAAAA 50A.ariommus B AAAAABAAAAAAA 1A.ariommus C AAAAAABAAAAAA 1A.constellatus ABAABABBABBBA 21M. salmoides BCABCACCBCCCB 1L. cyanellus CDACDCDDCBDDC 1population. Haplotype B had a unique restriction site for theendonuclease EcoRV, whereas haplotype C had a uniquerestriction site for Mlul. There was a single haplotype foundwithin the Ozark bass sampled (h =0.00). Unique restrictionprofiles between the Ozark and shadow basses wereidentified for the following restriction endonucleases: BgH,Dral, Pstl, Sail, Seal, and Xbal.Estimated nucleotide sequence divergence, standarderror, and time of divergence for Ambloplites, Lepomis, andMicropterus are found in Table 2. Divergence values rangedfrom 0.0612 for the shadow and Ozark basses to 0.3216 forthe largemouth bass and green sunfish. The divergencevalues for the shadow and Ozark basses are lower than thatbund for congeneric members within the genera Lepomisand Micropterus (Johnson and Williams,2003; Johnson et al.,2002). Data indicated that Ambloplites is more similar to theargemouth bass than to the green sunfish. Estimated timeofdivergence for the two species of Ambloplites studied was3.07 million years ago (Table 4). Unfortunately, there is noreported fossil record of Ambloplites on which to base acomparison. Itmust be noted that the molecular clock of2%)er million years has not been consistent through all taxaGrewe et al., 1990). Additionally, these estimates are basedon few populations within each species and therefore thevalues obtained are at best estimations of the divergence>eriods between these particular populations rather than thespecies as a whole.Construction of a cladogram generated utilizing the)ollo parsimony algorithm program revealed thatAmbloplites was most similar to Micropterus and less so toLepomis (Fig. 1). A UPGMA dendrogram derived from theaverage genetic distances between species was consistentwith that above. A cophenetic correlation of 0.76 wasTable 2. Mitochondrial DNA sequence divergence (abovediagonal) and estimated time of divergence in millions ofyears (below diagonal) for Ambloplites, Lepomis andMicropterus studied. Standard errors of the mean inparentheses beneath the means.Species 12 3 41. A.ariommus *** 0.0615 0.2044 0.1675***(0.0014) (0.0003) (0.0030)2. A. constellates 3.07 *** 0.2121 0.1371(0.01) ***(0.0004) (0.0006)3. L.cyanellus 10.22 10.66 *** 0.3216(0.01) (0.01) ***(0.0011)4. M.salmoides 8.37 6.86 16.08***(0.01) (0.01) (0.01) ***obtained, indicative of a suitable fit between the treegenerated and the data matrix (Rohlf, 1990). In contrast,Avise and Smith (1977) hypothesized that Lepomis wasintermediate to Micropterus and Ambloplites based uponallozyme analysis of several centrarchid genera. However,Avise and Smith (1977) sampled the rock bass as comparedto the shadow and Ozark basses, and they used the averagegenetic distances of ten Lepomis species rather than lookingat a specific species (green sunfish). Future study of thegenus Ambloplites at the population, subspecies, and specieslevels is required to further our understanding of thehistorical biogeography of this genus. Additional populationdata may alter the proposed relationships of these taxa.We would like to thank J. Quinn, K. Shirley, M. Wine,C. Hutts, T. Sanders and L. Mommsen for technicalassistance and for assistance in the collection ofspecimens.This work was supported by grants from the SiloUndergraduate Research Foundation and Arkansas StateUniversity.Literature CitedAvise, J. C, and M. H. Smith. 1977. Gene frequencycomparisons between sunfish (Centrarchidae)populations at various stages of evolutionarydivergence. Syst. Zool. 26: 319-335Branson, B. A., and G. A. Moore. 1962. The lateraliscomponents of the acoustico-lateralis system in thesunfish family Centrarchidae. Copeia 1962: 1-108.Journal of the Arkansas Academy of Science, Vol. 57, 2003207Journal of the Arkansas Academy of Science, Vol. 57 [2003], Art. 32http://scholarworks.uark.edu/jaas/vol57/iss1/32208Ronald L.Johnson and Carrie F. CavenaughL. cyanellusM.salmoidesA. constellatusA. ariommus 'CA.ariommus 'B'A.ariommus 'A'Fig. 1. Cladogram of character state matrices for presence/absence ofpresumptive restriction sites utilizing the Dolloparsimonyalgorithm. One parsimonious tree (22 steps) was generated.Brown, W. M.,M.George, Jr., and A. C. Wilson. 1979.Rapid evolution of animal mitochondrial DNA. Proc.Nat. Acad. Sci. 76:1967-1971.Cashner, R. C,and R.E.Jenkins. 1982. Systematics of theRoanoke bass, Ambloplites cavifrons. Copeia 1982:581-594.Felsenstein, J. 1993. Phylip (Phylogeny Inference Package).Version 3.51. University of Washington, Seattle.Grewe, P. M.,N. Billington, and P. D.N. Hebert 1990.Phylogenetic relationships among members ofSalvelinus inferred from mitochondrial DNAdivergence. Can. J. Fish. Aq.Sci. 47:984-991.Johnson, R. L., and K. A. Williams. 2003. Geneticrelationships of some common Arkansas freshwatersunfishes (Centrarchidae: Lepomis) inferred fromrestriction endonuclease analysis of mitochondrialDNA.J. Arkansas Acad. Sci. 57:(in press).Johnson, R. L., J. B. Magee, and T. A. Hodge. 2002.Phylogenetics of freshwater black basses(Centrarchidae: Micropterus) inferred from restrictionendonuclease analysis of mitochondrial DNA.Biochem. Gen. 39:395-496.Koppleman, J. B., C.M.Gale, andj. S. Stanovick. 2000.Allozyme and morphological variation among threenominal species of Ambloplites (Centrarchidae)inhabiting the Ozarks Region. Trans. Am. Fish. Soc.129:1134-1149.Nei, M., and W. H. Li. 1979. Mathematical models forstudying genetic variation in terms of restrictionendonucleases. Proc. Nat. Acad. Sci. 76:5269-5273.Nei, M., and F. Tajima. 1981. DNA polymorphismsdetectable by restriction endonucleases. Genetics97:147-165.Robison, H. W, and T. M. Buchanan. 1988. Fishes ofArkansas. University of Arkansas Press, Fayetteville.563 pp.Rohlf, F. F. 1990. NTSYS-PC: Numerical Taxonomy andMultivariate Analysis System. Version 1.60. ExeterPublishing, Setauket.Schaeffer, H. E., and R. R. Sederoff. 1981. Improvedestimation on DNA fragment lengths from agarose gels.Ann. Biochem. 115:113-122.Sokal, R. R., and P. H. A. Sneath. 1963. Principles ofnumerical taxonomy. Freeman, San Francisco. 359 pp.Journal of the Arkansas Academy of Science, Vol. 57, 2003208Journal of the Arkansas Academy of Science, Vol. 57 [2003], Art. 32Published by Arkansas Academy of Science, 2003

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Genetic Similarity of Shadow and Arkansas Basses (Ambloplites) Determined by Mitochondrial DNA Analysis

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