Sc.B./Sc.M.      (Appl. Math.)        Brown University         1974
M.S.             (Math.)              Stanford University      1976
Ph.D.            (Math.)              Stanford University      1979

Professional Experience
Assistant Professor             Purdue University             1979-1983
Assistant Professor             University of Northern Iowa   1983-1988
Associate Professor             University of Northern Iowa   1988-

Professional Societies and Honors
American Mathematical Society
American Society of Naturalists
Genetics Society of America
Mathematical Association of America
Sigma Xi
  Vice-President/President/Past President UNI Sigma Xi Club/Chapter 1988-1991
  (prepared successful petition for chapter status)
Society for the Study of Evolution

Research Interests
Mathematical Genetics
Evolution Theory

Theoretical Population Biology, Evolution, American Naturalist, Genetics, 
Biometrics, Pakistan Journal of Statistics, American Journal of
Human Genetics, Journal of Heredity, Journal of Cost Analysis and
Management, Mathematical Reviews, Journal of Theoretical Biology, National
Science Foundation, New Phytologist, Bulletin for Mathematical Biology, 
Journal of Mathematical Biology, Journal of Evolutionary Biology, McGraw Hill, 
Benjamin Cummings, Saunders, Duxbury, Wiley, Macmillan
Associate Editor: Theoretical Population Biology (1991-2009)

Numerous contributed papers have been presented at the Annual Meetings of The 
American Society of Naturalists, Genetics Society of America, and Society for 
the Study of Evolution; as well as a paper at the summer meeting of the AMS.  


1.  (with S. Karlin)  Analysis of Central Equilibrium Configurations for    
       Certain Multilocus Systems in Subdivided Populations.  Genetical
       Research (Cambridge) 32 (1978), 151-169.
2.  Polymorphic Equilibria with Assortative Mating and Selection in
       Subdivided Populations.  Theoretical Population Biology 18 (1980),
3.  (with S. Karlin)  Polymorphism in Subdivided Populations Characterized
       by a Major and Subordinate Demes.  Heredity 44 (1980), 151-168.
4.  The Effect of Migration and Recombination on the Equilibrium Structure
       of Populations Subject to a Common Symmetric Selection Regime. 
       Genetical Research (Cambridge) 36 (1980), 29-40.
5.  (with S. Karlin)  Selection-Migration Regimes Characterized by a
       Globally Stable Equilibrium.  Genetics 94 (1980), 1065-1084.
6.  (with S. Karlin)  The Existence of a Protected Polymorphism Under
       Conditions of Soft as Against Hard Selection in a Multideme
       Population System.  American Naturalist 117 (1981), 262-275.
7.  Some Circumstances Assuring Monomorphism is Sub-divided Populations. 
       Theoretical Population Biology 20 (1981), 118-125.
8.  The Effect of Variable Environments on Polymorphism at Loci with Several 
       Alleles I.  A Symmetric Model.  Journal of Mathematical Biology 13
       (1981), 199-208.
9.  Hard Selection in Haploid Species.  Theoretical Population Biology 21
       (1982), 1-10.
10. Note on Migration Modification.  American Naturalist 120 (1982), 
11. (with D. Hartl) Allele Multiplicity in Simple Mendelian Disorders. 
       American Journal of Human Genetics 34 (1982), 866-873.
12. The Effect of Variable Environments on Polymorphism at Loci with Several
       Alleles II.  Submultiplicative Viablilties.  Journal of Mathematical
       Biology 15 (1982), 293-303.
13. Repeatability of Experiments.  Bulletin of Mathematical Biology 44 (1982), 
14. Uniqueness of Polymorphic Equilibria under Soft Selection.  Theoretical
       Population Biology 24 (1983), 295-301.
15. The Manifestation of Phenotypic Selection at Constituent Loci.  I.
       Stabilizing Selection.  Evolution 38 (1984), 1033-1038.
16. Hercules' Height.  UMAP Journal 5 (1984), 265-269.
17. Dimension Reduction Projection and our Perception of Evolution.  Journal
       of Mathematical Biology 21 (1985), 299-306.
18. The Interdependence of Mating Structure and Inbreeding Depression. 
       Theoretical Population Biology 30 (1986), 232-244.
19. The Effects of Genetic Screening and Assortative Mating on Lethal
       Recessive-Allele Frequencies and Homozygote Incidence.  American
       Journal of Human Genetics 41 (1987), 671-677.
20. Mating Structure and the Cost of Deleterious Mutations.  I. The Effect
       of Postponing Inbreeding.  Journal of Heredity 79 (1988), 179-183.
21. Regular Systems of Inbreeding with Mutation.  Theoretical Population
       Biology 34 (1988), 24-37.
22. The Cumulation of Negligible Effects.  American Journal of Human
       Genetics 43 (1988),. 
23. Letter to the Editor.  Consortium 30 (1989), 2. 
24. On the Robustness of Regular Systems of Inbreeding.  Mathematical 
       Biosciences 104 (1991), 1-19. 
25. Rational Election Procedures. in J. G. Michaels and K. H. Rosen, eds., 
       Applications of Discrete Mathematics.  McGraw Hill.  (1991), 40-56.
26. The Apportionment Problem. in J. G. Michaels and K. H. Rosen, eds., 
       Applications of Discrete Mathematics.  McGraw Hill.  (1991), 2-18. 
27. Half-Sib Mating Structures.  Journal of Mathematical Biology 31 (1993), 
28. The Importance of Mating Structure versus Progeny Distribution for 
       Genetic Identity under Mutation.  Theoretical Population Biology 43 
       (1993), 129-140.
29. The effect of mating structure and progeny distribution on heterozygosity 
       versus number of alleles as measures of variation.  Journal of 
       Theoretical Biology 175 (1995), 503-509. 
30. The coalescent time in the presence of background fertility selection.   
       Theoretical Population Biology 55 (1999), 260-269.
31. Arbuthnot and the human sex ratio.  In: Evolution 2000, (ed. M. J. Wade),
       Indiana University Conferences. Bloomington, Indiana (2000).
32. Fertility selection, genetic selection, and evolution. Journal of  
       Evolutionary Biology 13 (2000), 786-791. 
33. John Graunt, John Arbuthnott, and the human Sex ratio.  Human Biology
       73 (2001), 605-610.
34. A constant regularity observ'd.  Math Horizons 11(1) (2003), 23-26.
35. A logistic branching process for population genetics.  Journal of 
       Theoretical Biology 225(2) (2003), 195-203.
36. Motivating mathematical concepts with politics. in J. L. Perry and S. G. 
       Jones, ed.,  Quick Hits for Educating Citizens. Indiana University Press.  
       (2006), 39-40.  
37. Coalescent size versus coalescent time with strong selection. Bulletin of 
       Mathematical Biology 69 (2007), 2249-2259.
38. Time since common pedigree ancestors with two progeny per individual. Mathematical  
       Population Studies 16(4) (2009), 248-265.
39. The Ancestry of Genetic Segments.  ISRN Biomathematics 2012. (Article ID 384275, 8 pages, 
40. First Day Statistics Activity - Grouping Qualitative Data.  STatistics Education Web 
      ( (2014)
41. Mating prescription, proscription, and the time since a common ancestor, a coalescent 
     approach. F1000Posters (; (2014). 
42. The effect of inbreeding constraints and offspring distribution on time to the most recent  
     common ancestor. Journal of Theoretical Biology 382 (2015), 74-80. 
     doi: 10.1016/j.jtbi.2015.06.037
43. The Impact of Lethal Recessive Alleles on Bottlenecks with Implications for Conservation
     Genetics.  BIORXIV/2016/089151
44. When did history occur?  Chance 32(4) (2019), 44-45.

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