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Systematics and the Origin of Species: On Ernst Mayr's 100th Anniversary (2005)
National Academy of Sciences (NAS)

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. "3 Inter-Locus Antagonistic Coevolution as an Engine of Speciation: Assessment with Hemiclonal Analysis--WILLIAM R. RICE, JODELL E. LINDER, URBAN FRIBERG, TIMOTHY A. LEW, EDWARD H. MORROW, AND ANDREW D. STEWART." Systematics and the Origin of Species: On Ernst Mayr's 100th Anniversary. Washington, DC: The National Academies Press, 2005.

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Systematics and The Origin of Species: On Ernst Mayr’s 100th Anniversary

fitness (P = 0.029), corroborating the significant positive net-selection gradient on this trait deduced from the measures of male adult lifetime offspring production from the larger sample of 35 hemiclones.

In sum, we found significant additive genetic variation among hemiclones for remating rate in both males and females, but the net-selection gradient on this trait was positive in males and negative in females. To look for independent genetic variation for remating rate in males and females, we constructed a bivariate plot of remating rate of the same hemiclones when expressed in males vs. females. No significant correlation was found (r = −0.175, P = 0.316), indicating that remating rate in the two sexes is controlled by different genetic variation. Because there is independent genetic variation for remating rate in the two sexes, and because it is selected in opposite directions in each sex, we conclude that this trait presently is evolving in opposite directions in the two sexes and therefore that sexually antagonistic coevolution for mating rate is currently in evidence in this laboratory island population.

INTERPRETATION OF RESULTS FROM HEMICLONAL ANALYSIS

In the above section, we used hemiclonal analysis to provide evidence that (i) females have genetic variation for resistance to male-induced harm, (ii) resistance contributes substantially to total genetic variation for net fitness, (iii) propensity to remating strongly influences the degree of female resistance, and (iv) there is unique genetic variation for remating rate in males and females that is selected and evolving in opposite directions in the two sexes. These data provide support for the hypothesis that perpetual inter-locus, intersexual arms races contribute to rapid genetic divergence among allopatric populations, and owing to the phenotypes that coevolve (reproductive behavior, physiology, and anatomy) are likely to be contributing to the specific genetic divergence that leads to reproductive isolation and speciation.

The data that we described, however, came from a laboratory island population rather than directly from nature. Some might argue that such populations are too artificial and hence tell us nothing about evolution in nature. We disagree. We cannot statistically extrapolate from our laboratory island population to natural populations of D. melanogaster because our laboratory population is not a random sample from the natural environment. We can, however, use laboratory island populations to make inferences about the fundamental principles of evolution and then use logic to extrapolate to the process of evolution in nature. Just as Darwin (1859) used his study of island tortoise populations to deduce general evolutionary principles (rather than extrapolate to specific continental

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Front Matter (R1-R14)
1 Introductory Essay: Systematics and the Future of Biology--EDWARD O. WILSON (1-4)
Part I--THE ORIGINS OF SPECIES BARRIERS: 2 The Genetic Basis of Reproductive Isolation: Insights from Drosophila--H. ALLEN ORR (5-23)
3 Inter-Locus Antagonistic Coevolution as an Engine of Speciation: Assessment with Hemiclonal Analysis--WILLIAM R. RICE, JODELL E. LINDER, URBAN FRIBERG, TIMOTHY A. LEW, EDWARD H. MORROW, AND ANDREW D. STEWART (24-45)
4 Chromosome Speciation: Humans, Drosophila, and Mosquitoes--FRANCISCO J. AYALA AND MARIO COLUZZI (46-68)
5 Developmental Plasticity and the Origin of Species Differences--MARY JANE WEST-EBERHARD (69-90)
Part II--DISCERNING RECENT DIVERGENCE: 6 Speciation in Birds: Genes, Geography, and Sexual Selection--SCOTT V. EDWARDS, SARAH B. KINGAN, JENNIFER D. CALKINS, CHRISTOPHER N. BALAKRISHNAN, W. BRYAN JENNINGS, WILLIE J. SWANSON, AND MICHAEL D. SORENSON (91-119)
7 Critical Review of Host Specificity and Its Coevolutionary Implications in the Fig/Fig-Wasp Mutualism--CARLOS A. MACHADO, NANCY ROBBINS, M. THOMAS P. GILBERT, AND EDWARD ALLEN HERRE (120-142)
8 Evolutionary Animation: How Do Molecular Phylogenies Compare to Mayr’s Reconstruction of Speciation Patterns in the Sea?--STEPHEN R. PALUMBI AND H. A. LESSIOS (143-161)
9 Mayr, Dobzhansky, and Bush and the Complexities of Sympatric Speciation in Rhagoletis--JEFFREY L. FEDER, XIANFA XIE, JUAN RULL, SEBASTIAN VELEZ, ANDREW FORBES, BRIAN LEUNG, HATTIE DAMBROSKI, KENNETH E. FILCHAK, AND MARTIN ALUJA (162-181)
10 On the Origin of Lake Malawi Cichlid Species: A Population Genetic Analysis of Divergence--YONG-JIN WON, ARJUN SIVASUNDAR, YONG WANG, AND JODY HEY (182-200)
Part III--THE NATURE OF SPECIES AND THE MEANING OF ‘‘SPECIES’’: 11 A Multidimensional Approach for Detecting Species Patterns in Malagasy Vertebrates--ANNE D. YODER, LINK E. OLSON, CAROL HANLEY, KELLIE L. HECKMAN, RODIN RASOLOARISON, AMY L. RUSSELL, JULIE RANIVO, VOAHANGY SOARIMALALA, K. PRAVEEN KARANTH, ACH (201-228)
12 Examining Bacterial Species Under the Specter of Gene Transfer and Exchange--HOWARD OCHMAN, EMMANUELLE LERAT, AND VINCENT DAUBIN (229-242)
13 Ernst Mayr and the Modern Concept of Species--KEVIN DE QUEIROZ (243-264)
Part IV--GENOMIC APPROACHES AND NEW INSIGHTS ON DIVERSITY: 14 Decoding the Genomic Tree of Life--ANNE B. SIMONSON, JACQUELINE A. SERVIN, RYAN G. SKOPHAMMER, CRAIG W. HERBOLD, MARIA C. RIVERA, AND JAMES A. LAKE (265-285)
15 Prospects for Identifying Functional Variation Across the Genome--STUART J. MACDONALD AND ANTHONY D. LONG (286-306)
16 Genetics and Genomics of Drosophila Mating Behavior--TRUDY F. C. MACKAY, STEFANIE L. HEINSOHN, RICHARD F. LYMAN, AMANDA J. MOEHRING, THEODORE J. MORGAN, AND STEPHANIE M. ROLLMANN (307-331)
17 Genomes, Phylogeny, and Evolutionary Systems Biology--MÓNICA MEDINA (332-350)
Index (351-368)