University of North Dakota UND Scholarly Commons Theses and Dissertations Theses, Dissertations, and Senior Projects 5-1994 Mitochondrial DNA Variation Within the Western-Clark's Grebe Complex (Podicipedidae: Aechmophorus) Bruce Alan Eichhorst Follow this and additional works at: https://commons.und.edu/theses Part of the Biology Commons Recommended Citation Eichhorst, Bruce Alan, "Mitochondrial DNA Variation Within the Western-Clark's Grebe Complex (Podicipedidae: Aechmophorus)" (1994). Theses and Dissertations. 3349. https://commons.und.edu/theses/3349 This Dissertation is brought to you for free and open access by the Theses, Dissertations, and Senior Projects at UND Scholarly Commons. It has been accepted for inclusion in Theses and Dissertations by an authorized administrator of UND Scholarly Commons. For more information, please contact [email protected]. MITOCHONDRIAL DNA VARIATION WITHIN' THE WESTERN-CLARK" S GREBE COMPLEX ( POOICIPBDIDAE :. ABCHMOPHORUS') by Bruce Alan Blchhor.st Bachelor of Science, Unive:rsi.t.y of Wisconsin-Oshkosh, 1982 Master of Sclence, Universi:ty of: Wisoonsin.-Oshkosh, 1986 A Di.ssertation. Submit.tad to the Graduate Faculty o.f the Uu:i vers.i'ty of North Dakota .ir. partlal. fulfillment, of 'the requirements for the de.gr.·e.e of Doct.<)r of: Philosophy Grand Forks, North Dakota May 1994 Copyrighted by Bruce Alan Eichhorst 1994 ii This dissertation, submitted by Bruce Alan Eichhorst in partial fulfillment of the x:equirements for the Degree of Doctor of Philosophy from the Univers.ity of North Dakota, has been read by the Faculty Advisory Committee under whom the work has been done and is hereby approved .. This dissertation meets the standards for appearance, conforms to the style and format requir·ements of t.he Gradua.te School of the Univer·sity of Mort.h Dakota, and is hereby approved .. _fl~ Dean of the G1du~ Scho~l t*~ i. ,,,, iii PERMISSION Title Mitochondrial DMA Variation Within the Western-Clark's Grebe Complex (Podicipedidae: Aechmophorus) Deportment Biology Degree Doctor of Philosophy In presenting this dissertation in partial fulfillment of the requirements for a graduate degree from the University of North Dakota, I agree that the library of this University shall make it freely available for inspection. I further agree that permission for extensive copying for scholarly purposes may be granted by the professor who supervised my dissertation work or, in his absence, by the chairperson of the department or the dean of the Graduate School. It is understood that any copying or publication or other use of this dissertation or part thereof for financial gain shall not be allowed without my written permission. It is also understood that due recognition shall be given me and to the University of North Dakota in any scholarly use which may be made of any material in my dissertation .. Signature ~(_4,~ Date ,;>. 7 ~ /7"71 iv TABLE OF CONTENTS LIST OF ILLUSTRATIONS .......................................... vii LIST OF TABLES., .................................................... ix ACKNOWLEDGME1'iTS ............................................xi ABSTAACT ••••••••••• " ..... ~ ................................ xiii CHAPTER I • GENERAL INTRODUCTION ................................. 1 II. BACKGROUND ON THE GENUS AECHMOPHORUS .••••.•..••..••• 4 Nomenclatural History ............................. 4 Fossil Histo·ry ••••••••••••• ,. ••••••••••.••••••••. 11 Current Distribution and Abundance.~·······~·~·-16 Migratory Movements •••••.••••••••••••••••...••••• 22 III. MITOCHONDRIAL DNA DIFFERENTIATION, POLYMORPHISM, AND POPULATION STRUCTURE IN THE WESTERN-CLARK'S GREBE COMPLEX (AECHMOPHORUS) OF THE UNITED STATES AND CANADA ................................................ 2 6 Introduction •.••••..•...••.••.••••..••.••••••••• 26 Materials and M~thods •••••• $ •••••••••••••••••••• 30 Choice of Sample Populations and Field Collections •••••••••••••••••••••••••••••••••• 30 Laboratory Analysis ••.•••••••.•.••••.••••••.• 34 Data .Analysis ................................. 35 Results ......................... ., •••••••••.••••• • 42 Mitochondrial-DNA Genome •••••••••••••••••••.• 42 Haplotype Phylogeny and Diversity .•••••.•.••• 44 Nucleotide Diversity and Divergence •..•••.••• 51 Population Genetic Variation ••.....•.. ~ •.•.•. 55 Discussion ••••.•••••.•..••.•.•••...••••.•••••••• 58 Phylogenetic Status of the Color Phases .••••• 58 Genetic Divergence Between the Color Phases,, .. 63 Intraspecific Phylogeography and Female­ mediated Gene Flow in Aechmophorus ..•.••••••• 68 Long-term Effective Population Size of Aechmophoz:us Females ............................ 7 0 Evolutionary and Taxonomic Implications •.•••• 72 V IV.. A REVIEW OF SPECIATION MODELS PROPOSED FOR THE WESTERN-CLARK'S GREBE COMPLEX (AECHMOPHORUS) •••.... 75 Introduction ••••..•••.•••.••..•••••••.••••••..•• 75 Quaternary History of Western North America •••.• 76 Aechmophorus Distribution and Migration ••••••••• 81 Aechmophorus Speciation Models •••••••••••••••••• 82 Discussion •.••••••••••••••••••••••••• o••••••••••84 Ratti's Models ••••• t••··~····················84 Feerer's Model ••••••••• ~·····················87 Conclusions •••.•.••••••••••••••••••••••••••••••• 88 V. RECOMMENDATIONS FOR FUTURE RESEARCH ON THE EVOLUTIONARY HISTORY OF THE AECHMOPBORUS GREBES •••• 89 APPENDIX ................................................... 9 3 LITERATURE CITED ........................................... 9 5 vi 1.. tocati OJfits wheilt.'e Ae:<ttlfimophx:,~utSJ f·.tf$:$ils ha,"\te be..e!t:l . dis'.00v·e:?ted.:. NUml:irelr:S· e:ort"ei$p0f.ld to ]oa:a.l.i.tie·s t.i:sJJed in, Te.:1b1l;e: 1 ~ .J .•: · .......... : ...•... ·• .......... •: l~: ~==:=::!!i::f}!f !ii~!!:![:liI~::ai:~:;/~~, ma. .:cs,;he.:s· wadthiili thf~'ise, ,crr:,ea1$.L.. N:umtJe~ .::;ifrewn. f;e>.~ e:a:eth st..artt.eilp.,f(l)v::i~ri'.e:-e, .i:s· .a ~·0X;tg1k, esAt±.ma:lt;e: ¢>:£' tttie.· fu±i!%!11,:::!::;:'.11:::~~P;Eii~;:::!;!::;! i;!::e'.de~::{t1& c·:~.1r1e.. nAt 1 e:he ~;te e,s;t.i'nra\t,eis: •. " !' .~· •· it: •. " i.• "!' .:, .•, ,ti; iii, ti) '!' .... ! • •' • •• • • if • • • i,, • • ~· • ··•·· ;j, ,/j,' ilii8i vii Figure Page 5. Unrooted parsimony network summar.1.z.1.ng restriction site changes between the 12 Aechmophorus m'LO-NA haplotypes. {a) Fragment patterns [A-CJ produced by the enzymes listed. Direction of arrow indicates inferred restriction site loss between adjacent fragment patterns. (b) Most parsimonious, unrooted network produced by both Dolle and Wagner parsimony methods, with the site changes indicated between adjacent haplotypes •••••• 38 6. UPGMA phenogram of the 12 Aechmophorus mtDNA haplotypes, based on genetic distances (d) between haplotypes ••••••••••••••••••••••••••••• 50 7. Map showing extent of Late Wisconsin glaciers in the Cascaae and Sierra Nevada ranges, and pluvial lakes within the Great Basin, western United States. Map was compiled and modified from maps in Porter et al. (1983) and Williams and Bedinger (1984)···~····························80 viii LIST OF TABLES Table Page 1. Aechmophorus fossil localities and the estimated ages of the fossil-bearing deposits. Distribution of localities shown in Figure 1 {identified by No.) ••••••••••••• ~··················14 2. Relative frequencies(%) of Clark's Grebes in various Aechmophorus breeding populations •••••••••• 19 3. Frequencies of mtDNA fragment patterns produced by restriction enzymes whose restriction sites were polymorphic within the total sample of 109 Aechmophorus grebes •••••••••••• 45 4. Distribution of rntDNA haplotypes within the Aechmophorus :;;,henotypes ............................. 46 5. Estimates of number of nucleotide substitutions per site (d) between 12 mtDNA haplotypes observed in 109 Aechmophorus grebes. d-values are above the diagonal, standard errors below the diagonal. All values expressed as percent. See Table 4 for genotypes ............................................ 4 7 6. Geographic distribution of Aechmophorus mtDNA hap 1 o type s • • • • • • • • • • • • • • • • • • • • • • • • • • .. • • • • • • • • • • • • • • 4 8 7. Estimates of mtDNA nucleotide (rr) and haplotypic (h) diversity in the Aechmophorus grebes surveyed in this study. rr expressed as percent ••...•••.••••••••••••••••••••••••••••••••••• 52 8. Average number of mtDNA nucleotide substitutions per site (dXY) and average number of net nucleotide substitutions per site(~) between Aechmophorus phenotypt.~s, and between geographic regions. All values expressed as percent ............................................. 5 3 ix Table Page 9. Average number of nucleotide substitutions per site ( dxn above diagonal; SE below diagonal) between Aechmophorus breeding sites. All values expressed as percent •••••••••••••••••••••••• 56 10. Average number of net nucleotide substitutions per site ( dAJ between breedj ,1g sites. All values expressed as percent •••••••••••••••••••••••• 57 11. Intrademe identity probabilities (underlined) for Aechmophorus breeding sites, and interdeme identity probabilities (below diagonal) between breeding sites ••••.•••• * ••••••••••••••••••••••••••• 58 12. Hierarchial analysis of molecular variance (AMOVA) on two matrices of mtDNA genetic distances between Aechmophorus haplotypes (see Materials and Methods for explanation). Only SW (breeding sites EAGL, GOOL, UKLL) and NE (ALKL, DELM, LOSA) geographic regions used in analysis •••••.••••••••••.•••••••••••••••••••••••••• 59 13. Estimated mtDNA genetic distances (percent nucleotide sequence divergence) reported for pairs of closely related