Download the full version of the textbook now at textbookfull.

com

Statistical Human Genetics Methods and

Protocols 2nd Edition Robert C. Elston (Eds.)

https://textbookfull.com/product/statistical-
human-genetics-methods-and-protocols-2nd-edition-
robert-c-elston-eds/

Explore and download more textbook at https://textbookfull.com

 Recommended digital products (PDF, EPUB, MOBI) that
you can download immediately if you are interested.

Reverse Genetics of RNA Viruses Methods and Protocols

Methods in Molecular Biology 2733 2nd Edition Daniel R.
Perez
https://textbookfull.com/product/reverse-genetics-of-rna-viruses-
methods-and-protocols-methods-in-molecular-biology-2733-2nd-edition-
daniel-r-perez/
textbookfull.com

Microbiome Analysis Methods and Protocols Robert G. Beiko

https://textbookfull.com/product/microbiome-analysis-methods-and-
protocols-robert-g-beiko/

textbookfull.com

Antibody Arrays: Methods and Protocols Kelly C. Whittaker

https://textbookfull.com/product/antibody-arrays-methods-and-
protocols-kelly-c-whittaker/

textbookfull.com

Online Banking Security Measures and Data Protection

Advances in Information Security Privacy and Ethics 1st
Edition Shadi A. Aljawarneh
https://textbookfull.com/product/online-banking-security-measures-and-
data-protection-advances-in-information-security-privacy-and-
ethics-1st-edition-shadi-a-aljawarneh/
textbookfull.com
Marriage Markets How Inequality is Remaking the American
Family 1st Edition June Carbone

https://textbookfull.com/product/marriage-markets-how-inequality-is-
remaking-the-american-family-1st-edition-june-carbone/

textbookfull.com

Strategic Innovation in Russia Towards a Sustainable and

Profitable National Innovation System 1st Edition Taco
C.R. Van Someren
https://textbookfull.com/product/strategic-innovation-in-russia-
towards-a-sustainable-and-profitable-national-innovation-system-1st-
edition-taco-c-r-van-someren/
textbookfull.com

Computer Vision in Control Systems-4: Real Life

Applications 1st Edition Margarita N. Favorskaya

https://textbookfull.com/product/computer-vision-in-control-
systems-4-real-life-applications-1st-edition-margarita-n-favorskaya/

textbookfull.com

Textbook of histology Leslie P. Gartner

https://textbookfull.com/product/textbook-of-histology-leslie-p-
gartner/

textbookfull.com

Priced Out The Economic and Ethical Costs of American

Health Care Uwe Reinhardt

https://textbookfull.com/product/priced-out-the-economic-and-ethical-
costs-of-american-health-care-uwe-reinhardt/

textbookfull.com
Market Momentum: Theory and Practice Stephen Satchell

https://textbookfull.com/product/market-momentum-theory-and-practice-
stephen-satchell/

textbookfull.com
Methods in
Molecular Biology 1666

Robert C. Elston Editor

Statistical
Human
Genetics
Methods and Protocols
Second Edition
METHODS IN MOLECULAR BIOLOGY

Series Editor
John M. Walker
School of Life and Medical Sciences
University of Hertfordshire
Hatfield, Hertfordshire, AL10 9AB, UK

For further volumes:

http://www.springer.com/series/7651
Statistical Human Genetics

Methods and Protocols

Second Edition

Edited by

Robert C. Elston
Case Western Reserve University, Cleveland, Ohio, USA
Editor
Robert C. Elston
Case Western Reserve University
Cleveland, Ohio, USA

ISSN 1064-3745 ISSN 1940-6029 (electronic)

Methods in Molecular Biology
ISBN 978-1-4939-7273-9 ISBN 978-1-4939-7274-6 (eBook)
DOI 10.1007/978-1-4939-7274-6
Library of Congress Control Number: 2017951222

© Springer Science+Business Media LLC 2017

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is
concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction
on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation,
computer software, or by similar or dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply,
even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations
and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to
be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty,
express or implied, with respect to the material contained herein or for any errors or omissions that may have been made.
The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Printed on acid-free paper

This Humana Press imprint is published by Springer Nature

The registered company is Springer Science+Business Media LLC
The registered company address is: 233 Spring Street, New York, NY 10013, U.S.A.
Preface

The recent advances in genetics, especially in the molecular techniques that have over the
last quarter of a century spectacularly reduced the cost of determining genetic markers, open
up a field of research that is becoming of increasing help in detecting, preventing, and/or
curing many diseases that afflict us. This has brought with it the need for novel methods of
statistical analysis and the implementation of these methods in a wide variety of computer
programs. The aim in this book is to make these methods and programs more easily
accessible to the beginner who has data to analyze, whether a student or a senior investiga-
tor. Twenty-eight of the thirty original chapters have been retained (all updated) and, in
view of changes in the field, three new chapters have been added. Apart from the first
chapter, which defines some of the genetic terms used in the book, each chapter of this book
takes up a particular topic and illustrates the use of at least one piece of software that the
authors have found helpful for the relevant statistical analysis of their own human genetic
data. There is often more than one program that performs a particular type of analysis and,
once you have used one program for a particular analysis, you may find you prefer another
program—and there is a good chance you will find that the same basic analysis is described in
more than one chapter of this book. You may therefore wish to browse over several chapters,
in the first place restricting your reading to only the introductory sections, which describe
the underlying theory. The chapters are ordered in the approximate logical order in which
human genetic studies are often conducted; so, if you are new to research in human genetics,
this initial reading could serve as an introduction to the subject. For the most part, the
second section of each chapter gives you step-by-step instructions for running programs and
interpreting the program outputs, with extra notes in the third section. However, although
the aim is very much to offer a “do it yourself” manual, there may well be times when you
will need to consult a statistical geneticist, especially for the interpretation of computer
output.

Cleveland, Ohio, USA Robert C. Elston

v
Contents

Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v
Contributors. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix
1 Statistical Genetic Terminology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Robert C. Elston, Jaya Satagopan, and Shuying Sun
2 Identification of Genotype Errors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Jeffery O’Connell and Yin Yao
3 Detecting Pedigree Relationship Errors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Lei Sun
4 Identifying Cryptic Relationships . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Lei Sun, Apostolos Dimitromanolakis, and Wei-Min Chen
5 Estimating Allele Frequencies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Indra Adrianto and Courtney Montgomery
6 Testing Departure from Hardy-Weinberg Proportions. . . . . . . . . . . . . . . . . . . . . . . 83
Jian Wang and Sanjay Shete
7 Estimating Disequilibrium Coefficients . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Maren Vens and Andreas Ziegler
8 Detecting Familial Aggregation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
Adam C. Naj and Terri H. Beaty
9 Estimating Heritability from Twin Studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
Katrina L. Grasby, Karin J.H. Verweij, Miriam A. Mosing,
Brendan P. Zietsch, and Sarah E. Medland
10 Estimating Heritability from Nuclear Family and Pedigree Data . . . . . . . . . . . . . . 195
Murielle Bochud
11 Correcting for Ascertainment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Warren Ewens and Robert C. Elston
12 Segregation Analysis Using the Unified Model . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
Xiangqing Sun
13 Design Considerations for Genetic Linkage and Association Studies . . . . . . . . . . 257
Jérémie Nsengimana and D. Timothy Bishop
14 Model-Based Linkage Analysis of a Quantitative Trait . . . . . . . . . . . . . . . . . . . . . . . 283
Yeunjoo E. Song, Sunah Song, and Audrey H. Schnell
15 Model-Based Linkage Analysis of a Binary Trait . . . . . . . . . . . . . . . . . . . . . . . . . . . . 311
Rita M. Cantor
16 Model-Free Linkage Analysis of a Quantitative Trait . . . . . . . . . . . . . . . . . . . . . . . . 327
Nathan J. Morris and Catherine M. Stein
17 Model-Free Linkage Analysis of a Binary Trait. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 343
Wei Xu, Jin Ma, Celia M.T. Greenwood, Andrew D. Paterson,
and Shelley B. Bull

vii
viii Contents

18 Single Marker Association Analysis for Unrelated Samples . . . . . . . . . . . . . . . . . . . 375

Gang Zheng, Ao Yuan, Qizhai Li, and Joseph L. Gastwirth
19 Single Marker Family-Based Association Analysis Conditional
on Parental Information . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 391
Ren-Hua Chung, Daniel D. Kinnamon, and Eden R. Martin
20 Single Marker Family-Based Association Analysis Not Conditional
on Parental Information . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 409
Junghyun Namkung and Sungho Won
21 Calibrating Population Stratification in Association Analysis. . . . . . . . . . . . . . . . . . 441
Huaizhen Qin and Xiaofeng Zhu
22 Cross-Phenotype Association Analysis Using Summary
Statistics from GWAS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 455
Xiaoyin Li and Xiaofeng Zhu
23 Haplotype Inference . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 469
Sunah Song, Xin Li, and Jing Li
24 Multi-SNP Haplotype Analysis Methods for Association Analysis . . . . . . . . . . . . . 485
Daniel O. Stram
25 The Analysis of Ethnic Mixtures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 505
Xiaofeng Zhu and Heming Wang
26 Detecting Multiethnic Rare Variants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 527
Weiwei Ouyang, Xiaofeng Zhu, and Huaizhen Qin
27 Identifying Gene Interaction Networks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 539
Danica Wiredja and Gurkan Bebek
28 Structural Equation Modeling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 557
Catherine M. Stein, Nathan J. Morris, Noémi B. Hall,
and Nora L. Nock
29 Mendelian Randomization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 581
Sandeep Grover, Fabiola Del Greco M., Catherine M. Stein,
and Andreas Ziegler
30 Preprocessing and Quality Control for Whole-Genome
Sequences from the Illumina HiSeq X Platform. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 629
Marvin N. Wright, Damian Gola, and Andreas Ziegler
31 Processing and Analyzing Human Microbiome Data . . . . . . . . . . . . . . . . . . . . . . . . 649
Xuan Zhu, Jian Wang, Cielito Reyes-Gibby, and Sanjay Shete

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 679
Contributors

INDRA ADRIANTO  Arthritis and Clinical Immunology Research Program, Oklahoma

Medical Research Foundation, Oklahoma City, OK, USA
TERRI H. BEATY  Department of Epidemiology, Johns Hopkins University Bloomberg School of
Public Health, Baltimore, MD, USA
GURKAN BEBEK  Systems Biology and Bioinformatics Graduate Program, Case Western
Reserve University School of Medicine, Cleveland, OH, USA; Center for Proteomics and
Bioinformatics, Case Western Reserve University School of Medicine, Cleveland, OH, USA;
Department of Nutrition, Case Western Reserve University School of Medicine, Cleveland,
OH, USA; Department of Electrical Engineering and Computer Science, Case Western
Reserve University School of Medicine, Cleveland, OH, USA
D. TIMOTHY BISHOP  Section of Epidemiology and Biostatistics, Leeds Institute of Cancer and
Pathology, University of Leeds, Leeds, UK
MURIELLE BOCHUD  Institute of Social and Preventive Medicine, Lausanne University
Hospital, Lausanne, Switzerland
SHELLEY B. BULL  Lunenfeld-Tanenbaum Research Institute, Sinai Health System, Toronto,
ON, Canada; Dalla Lana School of Public Health, University of Toronto, Toronto, ON,
Canada
RITA M. CANTOR  Department of Human Genetics, David Geffen School of Medicine at
UCLA, Los Angeles, CA, USA; Center for Neurobehavioral Genetics, Department of
Psychiatry, David Geffen School of Medicine at UCLA, Los Angeles, CA, USA
WEI-MIN CHEN  Center for Public Health Genomics, University of Virginia, Charlottesville,
VA, USA; Department of Public Health Sciences, University of Virginia, Charlottesville,
VA, USA
REN-HUA CHUNG  Division of Biostatistics and Bioinformatics, Institute of Population
Health Sciences, National Health Research Institutes, Miaoli County, Taiwan
FABIOLA DEL GRECO M.  Center for Biomedicine, EURAC Research, Bolzano, Italy
APOSTOLOS DIMITROMANOLAKIS  Department of Statistical Sciences, Faculty of Arts and
Science, Toronto, ON, Canada; Lunenfeld-Tanenbaum Research Institute, Mount Sinai
Hospital, Toronto, ON, Canada
ROBERT C. ELSTON  Case Western Reserve University, Cleveland, OH, USA
WARREN EWENS  Department of Biology, University of Pennsylvania, Philadelphia, PA, USA
JOSEPH L. GASTWIRTH  Department of Statistics, George Washington University,
Washington, DC, USA
DAMIAN GOLA  Institut f€ ur Medizinische Biometrie und Statistik, Universit€at zu L€ ubeck,
Universit€atsklinikum Schleswig-Holstein - Campus L€ ubeck, L€
ubeck, Germany
KATRINA L. GRASBY  Genetic Epidemiolgy, QIMR Berghofer Medical Research Institute,
Brisbane, Australia
CELIA M.T. GREENWOOD  Lady Davis Research Institute, Jewish General Hospital,
Montréal, QC, Canada; Department of Oncology and Department of Epidemiology,
Biostatistics & Occupational Health, McGill University, QC, Canada
SANDEEP GROVER  Institut f€ ur Medizinische Biometrie und Statistik, Universit€
a t zu L€
ubeck,
Universit€atsklinikum Schleswig-Holstein, L€
ubeck, Germany

ix
x Contributors

NOÉMI B. HALL  Department of Population and Quantitative Health Sciences, Case

Western Reserve University School of Medicine, Cleveland, OH, USA
DANIEL D. KINNAMON  Division of Human Genetics, Department of Internal Medicine, The
Ohio State University Wexner Medical Center, Columbus, OH, USA
QIZHAI LI  Academy of Mathematics and Systems Science, Chinese Academy of Sciences,
Beijing, China
XIAOYIN LI  Department of Population and Quantitative Health Sciences, School of
Medicine, Case Western Reserve University, Cleveland, OH, USA
XIN LI  Department of Electrical Engineering and Computer Science, Case Western Reserve
University, Cleveland, OH, USA
JING LI  Department of Electrical Engineering and Computer Science, Case Western Reserve
University, Cleveland, OH, USA
JIN MA  Lunenfeld-Tanenbaum Research Institute, Sinai Health System, Toronto, ON,
Canada
EDEN R. MARTIN  John P. Hussman Institute for Human Genomics, Leonard M. Miller
School of Medicine, University of Miami, Miami, FL, USA
SARAH E. MEDLAND  Genetic Epidemiolgy, QIMR Berghofer Medical Research Institute,
Brisbane, Australia
COURTNEY MONTGOMERY  Arthritis and Clinical Immunology Research Program,
Oklahoma Medical Research Foundation, Oklahoma City, OK, USA
NATHAN J. MORRIS  Department of Population and Quantitative Health Sciences, Case
Western Reserve University School of Medicine, Cleveland, OH, USA
MIRIAM A. MOSING  Department of Neuroscience, Karolinska Institute, Stockholm, Sweden;
Department of Medical Epidemiology and Biostatistics, Karolinska Institute, Stockholm,
Sweden
ADAM C. NAJ  Department of Biostatistics, Epidemiology, and Informatics, University of
Pennsylvania Perelman School of Medicine, Philadelphia, PA, USA; Department of
Pathology and Laboratory Medicine, University of Pennsylvania Perelman School of
Medicine, Philadelphia, PA, USA; Center for Clinical Epidemiology and Biostatistics,
University of Pennsylvania Perelman School of Medicine, Philadelphia, PA, USA
JUNGHYUN NAMKUNG  Molecular Diagnostics Team, IVD Business Unit, SK Telecom, Seoul,
South Korea
NORA L. NOCK  Department of Population and Quantitative Health Sciences, Case Western
Reserve University School of Medicine, Cleveland, OH, USA
JÉRÉMIE NSENGIMANA  Section of Epidemiology and Biostatistics, Leeds Institute of Cancer
and Pathology, University of Leeds, Leeds, UK
JEFFERY O’CONNELL  University of Maryland, Baltimore, MD, USA
WEIWEI OUYANG  Department of Global Biostatistics and Data Science, Tulane University
School of Public Health and Tropical Medicine, New Orleans, LA, USA
ANDREW D. PATERSON  Genetics and Genome Biology, The Hospital for Sick Children,
Toronto, ON, Canada; Dalla Lana School of Public Health, University of Toronto, Toronto,
ON, Canada
HUAIZHEN QIN  Department of Global Biostatistics and Data Science, Tulane University
School of Public Health and Tropical Medicine, New Orleans, LA, USA; Department of
Population and Quantitative Health Sciences, Case Western Reserve University School of
Medicine, Cleveland, OH, USA
CIELITO REYES-GIBBY  Department of Emergency Medicine, The University of Texas MD
Anderson Cancer Center, Houston, TX, USA
 Contributors xi

JAYA SATAGOPAN  Memorial Sloan Kettering Cancer Center, New York, NY, USA
AUDREY H. SCHNELL  Cardiovascular Imaging Core Laboratory, Harrington Heart &
Vascular Institute, University Hospitals Cleveland Medical Center, Cleveland, OH, USA
SANJAY SHETE  Department of Biostatistics, The University of Texas MD Anderson Cancer
Center, Houston, TX, USA; Department of Epidemiology, The University of Texas MD
Anderson Cancer Center, Houston, TX, USA
YEUNJOO E. SONG  Department of Population and Quantitative Health Sciences, Case
Western Reserve University, Cleveland, OH, USA
SUNAH SONG  Case Western Reserve University, Cleveland, OH, USA
CATHERINE M. STEIN  Center for Proteomics and Bioinformatics, Case Western Reserve
University, Cleveland, OH, USA
DANIEL O. STRAM  Department of Preventive Medicine, Keck School of Medicine, University
of Southern California, Los Angeles, CA, USA
SHUYING SUN  Texas State University, San Marcos, TX, USA
LEI SUN  Department of Statistical Sciences, Faculty of Arts and Science, University of
Toronto, Toronto, ON, Canada; Division of Biostatistics, Dalla Lana School of Public
Health, University of Toronto, Toronto, ON, Canada
XIANGQING SUN  Department of Population and Quantitative Health Sciences, Case
Western Reserve University School of Medicine, Cleveland, OH, USA
MAREN VENS  Institut f€ ur Medizinische Biometrie und Epidemiologie,
Universit€atsklinikum Hamburg-Eppendorf, Hamburg, Germany
KARIN J.H. VERWEIJ  Genetic Epidemiolgy, QIMR Berghofer Medical Research Institute,
Brisbane, Australia; Behavioural Science Institute, Radboud University, HR Nijmegen,
The Netherlands
JIAN WANG  Department of Biostatistics, University of Texas MD Anderson Cancer Center,
Houston, TX, USA
HEMING WANG  Department of Population and Quantitative Health Sciences, Case Western
Reserve University School of Medicine, Cleveland, OH, USA; Division of Sleep and
Circadian Disorders, Brigham and Women’s Hospital, Boston, MA, USA; Division of Sleep
Medicine, Harvard Medical School, Boston, MA, USA
DANICA WIREDJA  Systems Biology and Bioinformatics Graduate Program, Case Western
Reserve University School of Medicine, Cleveland, OH, USA; Center for Proteomics and
Bioinformatics, Case Western Reserve University School of Medicine, Cleveland, OH, USA;
Department of Nutrition, Case Western Reserve University School of Medicine, Cleveland,
OH, USA
SUNGHO WON  Department of Public Health Science, Graduate School of Public Health,
Seoul National University, Seoul, South Korea
MARVIN N. WRIGHT  Institut f€ ur Medizinische Biometrie und Statistik, Universit€
a t zu
L€ubeck, Universit€a tsklinikum Schleswig-Holstein - Campus L€
ubeck, L€ubeck, Germany
WEI XU  Department of Biostatistics, Princess Margaret Cancer Centre, University Health
Network, Toronto, ON, Canada; Dalla Lana School of Public Health, University of
Toronto, Toronto, ON, Canada
YIN YAO  Unit of Genomic Statistics, Intramural Research Program, National Institute of
Mental Health, Bethesda, MD, USA
AO YUAN  Department of Biostatistics, Bioinformatics and Biomathematics, Georgetown
University, Washington, DC, USA
XIAOFENG ZHU  Department of Population and Quantitative Health Sciences, Case Western
Reserve University School of Medicine, Cleveland, OH, USA
Visit https://textbookfull.com
now to explore a rich
collection of eBooks, textbook
and enjoy exciting offers!
xii Contributors

XUAN ZHU  Department of Biostatistics, The University of Texas MD Anderson Cancer

Center, Houston, TX, USA
ANDREAS ZIEGLER  Institut f€ur Medizinische Biometrie und Statistik & Zentrum f€ ur
klinische Studien, Universit€
at zu L€
ubeck, Universit€
atsklinikum Schleswig-Holstein,
L€ubeck, Germany
BRENDAN P. ZIETSCH  School of Psychology, University of Queensland, Brisbane, Australia
 Chapter 1

Statistical Genetic Terminology

Robert C. Elston, Jaya Satagopan, and Shuying Sun

Abstract
Common terms used in statistical genetics with multiple meanings are explained and the terminology used
in subsequent chapters is defined. Statistical human genetics has existed as a discipline for over a century,
and during that time the meanings of many of the terms used have evolved, largely driven by molecular
discoveries, to the point that molecular geneticists, statistical geneticists, and statisticians often have
difficulty understanding each other. It is therefore imperative, now that so much of molecular genetics is
becoming an in silico and statistical science, that we have a well-defined, common terminology.

Key words Gene, Allele, Locus, Site, Genotype, Phenotype, Dominant, Recessive, Codominant,
Additive, Phenoset, Diallelic, Multiallelic, Polyallelic, Monomorphic, Monoallelic, Polymorphism,
Mutation, Complex trait, Multifactorial, Polygenic, Monogenic, Mixed model, Transmission proba-
bility, Transition probability, Epistasis, Interaction, Pleiotropy, Quantitative trait locus, Probit, Logit,
Penetrance, Transformation, Scale of measurement, Identity by descent, Identity in state, Kinship
coefficient, Haplotype, Phase, Multilocus genotype, Allelic association, Linkage disequilibrium,
Gametic phase disequilibrium

In this introductory chapter we give the original meanings of

various genetic and statistical terms (which are found in the older
literature), together with some of the various meanings that are
sometimes ascribed to them today, and how the terms will be
defined in the following chapters. For simplicity, exceptions are
ignored and what is stated is usually, but not invariably, true.

1 Gene, Allele, Locus, Site

The concept of a gene (the word itself was introduced by Bateson)

is due to Mendel, who used the German word “Faktor,” or factor.
Mendel used the word in the same way that we might call “hot” and
“cold” factors, not in the way that we call “temperature” a factor. In
other words, his factor was the level of what statisticians now call a
factor. In the original terminology, still used by some population
geneticists, genes occur in pairs on homologous autosomal

Robert C. Elston (ed.), Statistical Human Genetics: Methods and Protocols, Methods in Molecular Biology, vol. 1666,
DOI 10.1007/978-1-4939-7274-6_1, © Springer Science+Business Media LLC 2017

1
2 Robert C. Elston et al.

chromosomes (the normal human genome comprises 22 pairs of

autosomal chromosomes and the two sex chromosomes X and Y).
In this terminology the four blood groups A, B, O, and AB (defined
in terms of agglutination reactions) are determined by three (allelic)
genes: A, B, and O. Nowadays molecular geneticists do not call
these three factors genes, but rather “alleles,” defined as “alterna-
tive forms” of a gene that can occur at the same locus, or place, in
the genome. Whereas Drosophila geneticists used to talk of two loci
for a gene, and human geneticists used to talk of two genes at a
locus, modern geneticists talk of “two alleles of a gene” or “two
alleles at a locus”; this last, which is nowadays so common, is the
terminology that will thus be used in this book. It then follows
(rather awkwardly) that two alleles at the same locus are allelic to
each other, whereas two alleles that are at different loci are nonalle-
lic to each other. A gene is commonly defined as a DNA sequence
that has a function, meaning a class of similar DNA sequences all
involved in the same particular molecular function, such as the
formation of the ABO red cell antigens. (Note the common illogi-
cal use of the phrase “cloning genes” by molecular geneticists
when, by their own terminology, “cloning alleles” is meant).
Some restrict the word gene to protein-coding genes, but there
are many more sequences of DNA that have function by virtue of
being transcribed to RNA without ever being translated to DNA
and then coding for a protein, so this restricted definition of a gene
would appear to be unwarranted. See [1] for a more detailed
explanation of the evolution of, and a modern definition of, the
word “gene.”
A locus is the location on the genome of a gene, such as the
“ABO gene.” By any definition a gene must involve more than one
nucleotide base pair. Single nucleotide polymorphisms (SNPs) thus
do not occur at loci, but rather in and around loci, and in this book
we shall not write of SNP markers as being “at” loci. Because of the
confusion that occurs when SNPs are described as occurring at loci,
some use the term “gene-locus,” but we shall always use the term
locus for the location of a functional gene, whether or not it is
protein coding. We shall, however, allow SNP markers to have
alleles and use the original term if necessary for their locations:
“sites” within loci or, more generally, sites within the region of a
locus or anywhere in the genome.
If in the population only one allele occurs at a site or locus, we
shall say that it is monomorphic, or monoallelic, in that population.
If two alleles occur, as is common for nonmonomorphic SNPs, we
shall use the original term diallelic which, apart from having prece-
dence, is etymologically sounder than the now commonly used
term biallelic. If many alleles occur, we shall describe the polymor-
phism as polyallelic or multiallelic (the former term is arguably
more logical, the latter more common). When there are just two
alleles at a locus or site, the one with the smaller population
 Statistical Genetic Terminology 3

frequency is called the minor allele. In genetics, in the context of

alleles the term “frequency”—which in statistics is a count—is used
to mean relative frequency, i.e., the proportion of all such alleles at
that locus among the members of a population; thus the term
minor allele frequency is often used to indicate the less frequent
allele in the case of a diallelic SNP.

2 Genotype, Phenotype, Dominant, Recessive, Codominant, Additive

An individual’s genotype is the totality of that individual’s heredi-

tary material, whereas an individual’s phenotype is the individual’s
observed appearance. However, the terms genotype and phenotype
are usually used in reference to a particular locus or set of loci, and
to a particular trait or set of traits. Genotypes are not observed
directly, but rather inferred from particular phenotypes. Thus, with
respect to the ABO locus, the four blood types A, B, O, and AB are
(discrete) phenotypes; and the possible genotypes, formed by pairs
of alleles, are AA, AO, BB, BO, AB, and OO. With respect to the
ABO blood group phenotypes, the A allele is dominant to the O
allele and the O allele is recessive to the A allele. Similarly, the B
allele is dominant to the O allele and the O allele is recessive to the
B allele. The A and B alleles are codominant. Note that for the
words “dominant” and “recessive” to have any meaning, at least
two alleles and two phenotypes must be specified. If a particular
allele at a locus is dominant with respect to the presence of a disease,
there must be at least one other allele at that locus that is recessive
with respect to absence of that disease.
Geneticists loosely talk about a disease being dominant, mean-
ing that, with respect to the phenotype “disease,” the underlying
disease allele is dominant, i.e., the disease is present when either a
single or two copies of the allele is present. Similarly, they talk of a
disease being recessive, meaning that, with respect to the same
phenotype, the underlying disease allele is recessive, i.e., the disease
is present only when two disease alleles are present. Alternatively,
they may talk of an allele being dominant or recessive, the particular
phenotype (often disease) being understood. The important thing
to realize is that “dominance” and “recessivity” describe a relation-
ship between one or more genotypes and the distribution of a
phenotype. This leads to the concept of phenosets: the genotypes
AA and AO form the phenoset corresponding to the A blood type,
and the genotypes BB and BO form the phenoset corresponding to
the B blood type. In the case of the ABO blood group, a person
who has one A allele and one B allele has the blood type AB, which
is a phenotypically different from either of the corresponding
homozygotes, AA and BB; this relationship is called codominance.
In general, a locus is codominant with respect to the set of pheno-
types it controls if the phenotypes of each heterozygote at that
4 Robert C. Elston et al.

locus differ from that of each of the two corresponding homozy-

gotes. We make a distinction between codominant and additive; the
latter implies that the phenotype (or phenotypic distribution, see
below under quantitative phenotypes) corresponding to the het-
erozygote is in some sense halfway between those of the two
corresponding homozygotes. Whereas the term additive is only
meaningful when a scale of measurement has been defined, codo-
minance is a more general concept that does not require the defini-
tion of a scale of measurement.

3 Polymorphism, Mutation

The A, B, O, and AB blood types comprise a polymorphism, in the

sense that they are alternative phenotypes that commonly occur in
the population. A polymorphic locus was originally defined opera-
tionally as a polymorphism-determining locus at which the least
common allele occurs with a “frequency” of at least 1% [2]; but a
more appropriate definition would be a locus at which the most
common allele occurs with a “frequency” of at most 99%. Different
alleles arise at a locus as a result of mutation, or sudden change in
the genetic material. Mutation is a relatively rare event, caused for
example by an error in replication. Thus all alleles are by origin
mutant alleles, and a genetic polymorphism was conceived of as a
locus at which the frequency of the least common allele has a
frequency too large to be maintained in the population solely by
recurrent mutation. However, what is important at a locus is the
degree of polymorphism, and a locus in which there are 1000
equifrequent alleles would be considered much more polymorphic
than a locus at which there are two alleles with respective frequen-
cies 0.01 and 0.99. Many authors use the term mutation for any
rare allele, and the term polymorphism for any common allele. We
shall avoid this usage here.

4 Complex Trait, Multifactorial, Polygenic, Monogenic

The term “complex trait” was introduced about three decades ago
without a clear definition. It appears to be used for traits that do not
exhibit clear one-locus (“Mendelian”) segregation, usually because
segregation at more than one locus is involved. Whereas multifac-
torial and complex are often used interchangeably, a clear distinc-
tion should be made between the terms multifactorial and
polygenic. Multifactorial implies that more than one factor is
involved in the etiology of the phenotype, whether genetic, envi-
ronmental, or both. Polygenic, on the other hand, implies that only
genetic factors are involved, usually in an additive fashion, with the
original definition that the number of factors (loci) is so large that
 Statistical Genetic Terminology 5

they cannot be individually characterized. Thus, strictly speaking,

the term polygenic should not be used to include any environmen-
tal factors—though in practice it often is used that way.
Monogenic inheritance implies segregation at a single locus,
while polygenic inheritance implies the segregation of many loci.
The term “mixed model” is used by geneticists to denote an
additive combination of monogenic and polygenic inheritance. In
a segregation model, both components may be latent variables, i.e.,
unobserved variables (the former discrete and the latter continu-
ous). In this situation the underlying statistical model is random,
not mixed, because there are two random components other than
any error term. Confusion can also occur when statistical geneti-
cists use the term “transmission probability” in two quite different
senses. In this book we carefully distinguish transmission probabil-
ities, probabilities that a parent having a particular genotype trans-
mits particular alleles to offspring, from transition probabilities,
probabilities that offspring receive particular genotypes from their
parents. This distinction was introduced in [3].

5 Haplotype, Phase, Multilocus Genotype

Let A,B be two alleles at one locus, and D,d be two alleles at
another locus. If one parent transmits A and D to an offspring,
while the other transmits B and d, the offspring genotype is
denoted AD/Bd (or Bd/AD), in which the parental origins are
separated by “/”. The two alleles transmitted by one parent consti-
tute a two-locus haplotype. With respect to two alleles at each of
two loci there are four possible haplotypes—AD, Ad, BD, and Bd in
this case, with AD/Bd and Bd/AD being the two possible phases of
the two-locus genotypes. If n1 alleles can occur at one of the
loci and n2 at the other, n1n2 two-locus haplotypes are possible.
At the first locus n1(n1 þ 1)/2 genotypes are possible (n1 homo-
zygotes and n1(n1–1)/2 heterozygotes), while at the second locus
n2(n2 þ 1)/2 genotypes are possible. If we pair these genotypes,
one from each locus, the total number of pairs possible is
½n1 ðn1 þ 1Þ=2½n2 ðn2 þ 1Þ=2 ¼ n1 n2 ðn1 n2 þ 1Þ=2  ½n1 ðn1  1Þ=2
½ n2 ðn2  1Þ=2:
On the other hand, at the two loci together, there are n1n2
haplotypes; and pairing these we have n1n2(n1n2 þ 1)/2 possible
pairs of two-locus haplotypes, or diplotypes. In this book we shall
define “two-locus genotypes” this way, i.e., without differentiating
the two phases, so that for the same number of alleles at each locus
there is a smaller number of two-locus genotypes than there are
two-locus diplotypes. Thus we shall consider the two phases of
the double heterozygote, Ad/BD and BD/Ad, as being the same
6 Robert C. Elston et al.

two-locus genotype. Usually, the term “multilocus genotype”

refers to genotypes where the phases are not distinguished, and
the term diplotype is useful for the case when they are distinguished
(though this term is not yet in common usage).
More generally, a haplotype is the multilocus analogue of an
allele at a single locus. It consists of one allele from each of multiple
loci that are transmitted together from a parent to an offspring.
When haplotypes made up of multiple alleles (one from each locus)
are paired, a pair in which the genotype at each of n loci is hetero-
zygous corresponds to 2n–1 different diplotypes, or phases. It is
usual nowadays to restrict the word haplotype to the case where all
the loci involved are on the same chromosome pair, so that all the
alleles involved are on the same chromosome. Typically, but not
always, it is assumed that all the different phases of a particular
multiple heterozygote have the same phenotype.

6 Epistasis, Pleiotropy

When two loci are segregating, each typically influences a separate

phenotype. For example, A and B may be alleles at the ABO locus,
determining the ABO blood types, while D and d are alleles at a
disease locus, determining disease status. But if alleles at a single
locus influence two different phenotypes, we say there is pleiotropy.
It is known that a person’s ABO genotype influences the risk of
gastric cancer as well as determining blood type. Thus the ABO
locus is pleiotropic. Alternatively, alleles at two different loci may
determine the same phenotype, such as presence or absence of a
disease; and if the phenotype associated with the genotypes at one
locus depends on the genotypes at another locus, we say there is
epistasis. Thus gastric cancer may perhaps be caused by the epistatic
effect of alleles at two (or more) loci. Epistasis and pleiotropy are
sometimes confused in statistical genetics.

7 Allelic Association, Linkage Disequilibrium, Gametic Phase Disequilibrium

If the alleles at one locus are not distributed in the population

independently of the alleles at another locus, the two loci exhibit
allelic association. If this association is a result of a mixture of
subpopulations (such as ethnicities or religious groups) within
each of which there is random mating, the association is often
denoted as “spurious.” In such a case there is true association,
but the cause is not of primary genetic interest. If the association
is not due to this kind of population structure, it is either due to
linkage disequilibrium (LD) or gametic phase disequilibrium
(GPD); in the former case the loci are linked, i.e., they cosegregate
 Statistical Genetic Terminology 7

in families, in the latter case they are not linked, i.e., they segregate
independently in families. Owing to an unintended original defini-
tion, loci that are not linked have often been mistakenly described
as being in LD [4, 5].

8 Identity

The concept of allelic identity is an important one. Alleles are

identical by descent (IBD) if they are copies of the same ancestral
allele, and must be differentiated from alleles that are physically
identical but not (at least within the previous dozen or so genera-
tions) ancestrally identical. Such alleles, when not IBD, are identical
in state (IIS). It is well understood that molecules, atoms, etc., can
be in different states (not “by” different states), and the same is true
of alleles, though here the states are ancestrally, not physically,
different. Whereas in the animal and plant genetics literature
the phrases “identity in state” and “identical in state” are com-
monly used, for no good reason the phrases “identity by state” and
“identical by state” are now commonly used in the human genetics
literature. In this book, to stress the difference and to be consistent
with both the earlier common usage and the usage in the
animal and plant genetics literature, we shall use the terminology
IIS, not IBS.

9 Kinship Coefficient

The kinship coefficient between two individuals is a measure of

their genetic relationship on the assumption of additive inheritance.
Formally, the kinship coefficient between two individuals i and j is
the probability that a random allele at a given locus of i is identical
by descent with a random allele at the same locus of j. However, this
definition assumes the loci are autosomal, i.e., not on the sex
chromosomes X and Y. Furthermore, this definition ignores the
possibility of copy number variants, i.e., the possibility that an
individual has other than two alleles at a locus.

10 Quantitative Traits

A locus at which alleles determine the level of a quantitative pheno-

type is called a QTL (quantitative trait locus). Typically, the word
“quantitative” is used interchangeably with “continuous” when
describing a phenotypic distribution. However, quantitative traits
can be discrete (e.g., number of digits). Care should be taken to
distinguish between those methods of analysis of quantitative
traits for which distributional assumptions, such as conditional nor-
mality, are critical, and those for which they are not. Transforming the
8 Robert C. Elston et al.

phenotype of a QTL corresponds to changing its units if the trans-

formation is linear, or more generally to changing the scale of
measurement (e.g., square root or logarithmic) if the transforma-
tion is nonlinear. On the scale of measurement used, alleles at a QTL
have an additive effect if the phenotypic distribution of the hetero-
zygote is the average of the corresponding two homozygote pheno-
typic distributions. With respect to that phenotype, allele A is
dominant to the allele B, and allele B is recessive to allele A, if the
whole phenotypic distribution of the heterozygote AB is the same as
that of the homozygote AA. Any variance among the phenotypic
means of the genotypes at a locus over and above that due to
additive allele action is called dominance, or dominant genetic,
variance. Thus dominance variance can arise as a result of one allele
being dominant to another, but such simple allele action is not
necessarily implied by dominance variance. The presence of domi-
nance variance depends on the scale of measurement; dominant
allele action (complete dominance, as described above for discrete
traits such as the ABO blood group) does not. If the phenotypic
distribution of a heterozygote is not the average of the
corresponding two homozygote phenotypic distributions, we shall
say there is codominance. Thus, in this book, we shall not restrict the
word codominance to the case of additivity (which would result in
codominance being scale independent).
Just as dominance has a different meaning when applied to
quantitative traits, so does epistasis. From a statistical point of
view, dominance can be considered as intralocus interaction, or
non-additivity of the allelic contributions to the phenotype. Epista-
sis is a genetic term, now generalized when applied to quantitative
traits to indicate nonadditivity of the effects on the phenotype of
the genotypes at two (or more) loci in a population. It is thus from a
statistical viewpoint interlocus interaction, and so dependent on
how the phenotype is measured. Statistical interaction is a term
with a similar limitation, but is not restricted to genetic factors.
Statistical interaction should be carefully distinguished from
biological interaction [5, 6]. Whereas biological interaction does
not require the presence of statistical interaction, the presence of
the latter implies the existence of the former. Indeed, statistical
interaction, but not biological interaction, is removable if a mono-
tonic transformation can make the effects of the two factors
involved (e.g., segregation at two loci, or segregation at one locus
and levels of an environmental factor) additive. Furthermore, the
magnitude of any interaction effects can depend critically on how
the individual factor effects (single locus genotypes in the case of
genetic factors) are defined [5].
There is usually no loss of generality in assuming that disease
status, unaffected or affected, is a quantitative trait that takes on the
values 0 or 1, respectively, so that its mean value is the population
prevalence of the disease. Then everything that holds with regard to
dominant allele action, dominance variance and epistasis also holds
 Statistical Genetic Terminology 9

in the case of a binary disease phenotype, except that the scale of

measurement (in the sense of a nonlinear monotonic transforma-
tion) is irrelevant in the absence of a nonbinary quantitative mea-
sure. However, if there is such a quantitative measure being
analyzed, for example a relative risk or odds ratio, then the scale
of measurement will determine whether or not there is statistical
interaction. In the case of a binary disease phenotype the pene-
trance, or probability of being affected, is often transformed to a
probit (or logit), giving rise to what is called the “liability” to
disease, and this liability is treated as a continuous phenotype.
Note that it is the penetrance of the phenotype, not the phenotype
itself, that is transformed and analyzed as though it is a quantitative
trait. Dominance and epistatic variance can be quite different on
this liability scale from that measured on the original “penetrance”
scale.
For a QTL, dominance variance is present when there is intra-
locus nonadditivity. By the same token, epistatic variance is present
when there is interlocus nonadditivity. Each locus gives rise to its
own components of additive genetic and dominant genetic vari-
ance. If multiple loci affect a QTL, there are multiple components
of epistatic variance. Except in the case of a binary disease pheno-
type assigned distinct quantitative values to affected and unaffected
status, the relative magnitudes of all such components are scale (i.e.,
transformation) dependent, just as corresponding components of
genotype (or allele)  environment interaction are scale dependent.
Finally, for those who wish to have a better theoretical under-
standing of statistical human genetics, reference [7] provides an
exceptionally good introduction.

Acknowledgments

This work was supported in part by the following grants from the
National Institutes of Health, USA: P41RR003655 (RCE) and
R01CA137420 (JMS).

References
1. Gerstein MB et al (2007) What is a gene, post- 5. Wang X, Elston RC, Zhu X (2010) The meaning
ENCODE? History and updated definition. of interaction. Hum Hered 70:269–277
Genome Res 17:669–681 6. Wang X, Elston R, Zhu X (2010) Statistical
2. Ford EB (1940) Polymorphism and taxonomy. interaction in human genetics: how should we
In: Huxley J (ed) The new systematics, Oxford model it if we are looking for biological interac-
University Press tion? Nat Rev Genet 12:74. doi:10.1038/
3. Elston RC, Stewart J (1971) A general model for nrg2579-c2
the genetic analysis of pedigree data. Hum 7. Ziegler A, König IR (2010) A statistical
Hered 21:523–542 approach to genetic epidemiology: concepts
4. Lewontin RC (1964) The interaction of selec- and applications, 2nd edn. Wiley-VCH,
tion and linkage. I. General considerations; het- Weinheim
erotic models. Genetics 49:49–67
Visit https://textbookfull.com
now to explore a rich
collection of eBooks, textbook
and enjoy exciting offers!
 Chapter 2

Identification of Genotype Errors

Jeffery O’Connell and Yin Yao

Abstract
It has been documented that there exist some errors in most large genotype datasets and that an error rate of
1–2% is sufficient to lead to the distortion of map distance as well as a false conclusion of linkage (Abecasis
et al., Eur J Hum Genet 9:130–134, 2001), therefore one needs to ensure that the data are as clean as
possible. On the other hand, the process of data cleaning is tedious and demands effort and experience.
O’Connell and Weeks implemented four error-checking algorithms in computer software called PedCheck.
In this chapter, the four algorithms implemented in PedCheck are discussed with a focus on the genotype-
elimination method. Furthermore, an example for four levels of error checking permitted by PedCheck is
provided with the required input files. In addition, alternative algorithms implemented in other statistical
computing programs are also briefly discussed.

Key words Genotype, Genotype error, Parametric linkage analysis, LOD score, Computational
efficiency, Automatic genotype elimination, Nuclear pedigree method, Genotype-elimination method,
Critical genotype method, Odds ratio method

1 Introduction

While gene hunters have limited access to computational resources,

they have to rely on visual inspection to check for genotypic errors
occurring in a human pedigree. The errors come from two main
major sources: (1) pedigree errors and (2) true genotyping errors.
Pedigree errors include, but are not limited to, nonpaternity, unre-
ported adoption status, errors in data entry as well as sample mix-
ups. In this chapter, we focus only on true genotyping errors.
One can imagine that if automated approaches for error check-
ing were not available, detection of erroneous genotypes by visual
inspection could be a tedious task, particularly in extended pedi-
grees with multiple generations. Therefore, there is a demand for
computational algorithms that can efficiently identify erroneous
genotypes regardless the size of the pedigrees. Needless to say, the
elimination of genotype errors will benefit both linkage analysis and
family-based association analysis. In this chapter, we use linkage
analysis as an example for the sake of illustration.

Robert C. Elston (ed.), Statistical Human Genetics: Methods and Protocols, Methods in Molecular Biology, vol. 1666,
DOI 10.1007/978-1-4939-7274-6_2, © Springer Science+Business Media LLC 2017

11
12 Jeffery O’Connell and Yin Yao

Linkage analysis is a statistical method that has been widely

applied for mapping genes related to human disorders with rela-
tively high penetrance and rare disease allele frequencies. Because of
the fact that DNA segments located near each other on a chromo-
some tend to be passed together from one generation to another,
genetic markers are often used as tools for tracking the inheritance
pattern of a gene that has not yet been identified but the approxi-
mate location of the gene of interest is known.
The main use of linkage analysis is to test for evidence of linkage
between a disease locus of interest and an arbitrary marker locus.
The LOD score was defined as a test statistic by Morton [1]. A two-
point LOD score test is typically performed by maximizing the
LOD score over a grid of values of the recombination fraction θ
in the interval 0–0.5, where this fraction is the probability of
recombination, viewed as a measure of the extent of linkage. The
recombination fraction ranges from θ ¼ 0 for loci in close proximity
to each other through θ ¼ 0.5 for loci that are far apart or on
different chromosomes.
The formula for a LOD score is:

Z ðθÞ ¼ log10 ½L ðθÞ  log10 ½L ðθÞ ¼ 0:50:

In practice, LOD scores can be calculated using linkage analysis
programs such as LIPED [2] and LINKAGE [3–5]. A well-known
algorithm for parametric linkage analysis is the Elston–Stewart
algorithm [6], in which the genetic mode of inheritance is assumed,
and this algorithm was implemented in the LINKAGE software in
an elegant manner. It is common knowledge that LINKAGE is
capable of handling relatively large pedigrees, but the computation
time of LINKAGE is exponential in the number of genetic markers
[7]. In the presence of genotypic errors, the increase in computa-
tional time can be tremendous. Therefore, eliminating Mendelian
inconsistencies in pedigree data is an important task if only for the
sake of improving the efficiency of linkage analysis.
Lange and Goradia [8] described an algorithm for automatic
genotype elimination which enables great reduction in computa-
tional times required for pedigree analysis. Their genotype elimina-
tion program was an extension of the one given by Lange and
Boehnke [9], which is discussed in this chapter. Later, Stringham
and Boehnke [10] developed two methods that were implemented
in the Mendel computer software [11] to calculate the posterior
probability of erroneous genotypes for each pedigree member.
Stringham and Boehnke developed two novel approaches to com-
pute an individual posterior probability of genotype error using a
weighted sum of all possible genotypes, where the weight is the
probability of the genotype being an error. Their first approach
allows each individual to have every possible genotype weighted
 Identification of Genotype Errors 13

by the probability that the genotype is erroneous and then

computes the posterior probability of this individual’s genotype,
whereas their second approach allows one genotyped pedigree
member at a time to have every possible genotype while the other
pedigree members are held at their originally assigned genotypes.
They then compute the posterior probability for each genotyped
individual, based on the assumption that all other pedigree mem-
bers have been correctly genotyped. It has been recognized that
both methods have the weakness of not being able to automatically
deal with more than one marker at a time; therefore, the computa-
tion for markers with multiple alleles (e.g., microsatellites) is very
time consuming. These two algorithms were implemented in
MENDEL [11]. Of note, Sobel and Lange [12] developed impor-
tant extensions to the Lander–Green–Kruglyak algorithm for small
pedigrees and to the Markov-chain Monte Carlo stochastic algo-
rithm for large pedigrees to conduct linkage analysis while accom-
modating genotype errors. These developments have been
implemented in the program MENDEL, version 4.
In recognition of the need for developing efficient computer
software for automatic error checking, O’Connell and Weeks [13]
implemented four error-checking algorithms in a new computer
program called PedCheck. The main goal of this chapter is to
illustrate the algorithms used in PedCheck and, for the purpose of
demonstration, offer a few examples for different levels of error
checking permitted by PedCheck, as well as provide detailed
instructions for users to follow using the simulated pedigree data
presented in Figs. 1 and 2.

Fig. 1 Example of level one error

14 Jeffery O’Connell and Yin Yao

Fig. 2 Example of Mendelian error in a pedigree with three generations

1.1 The Nuclear Lange and Goradia [8] described the steps involved in a using
Pedigree Algorithm nuclear pedigree as follows:
A. For each pedigree member, save only those genotypes compat-
ible with his or her phenotype.
B. For each nuclear family:
1. Examine each mother-father genotype pair.
(a) Determine which zygote genotypes can arise.
(b) If each child in the nuclear family has one or more of
these zygote genotypes among his or her list of geno-
types, then save the parental genotypes as well as any
child genotype that is not conflicted with one of the
created zygote genotypes.
(c) If any child is incompatible with the current parental
pair, then take no action to save any genotypes.
2. For each person in the nuclear family, exclude any genotypes
not saved during step 1.
C. Repeat step B until no more genotypes can be excluded.

1.2 The Genotype- The genotype-elimination algorithm used by O’Connell and Weeks
Elimination Algorithm was an extension of the Lange–Goradia algorithm [8, 14] for set-
recoded genotypes [13]. Because the genotype-elimination algo-
rithm is able to detect subtle levels of inconsistencies due to the
elimination of certain genotypes presented in pedigrees with more
complex structure, it is more powerful than the nuclear-family
algorithm originally described by Lange and Goradia [8].
 Identification of Genotype Errors 15

For example, Fig. 1 shows that individual 7 cannot possibly carry

allele 3 given that the genotype of his father is 4/3 and of his
mother is 4/6.
As described by O’Connell and Weeks [13], for each pedigree
and locus, the genotype-elimination algorithm can pick the first
component nuclear family with an error that has been missed by the
nuclear family algorithm, and then provide the inferred-genotype
lists for each member of that nuclear family. To illustrate the situa-
tion where the nuclear-family algorithm is not able to find any
errors, we present an example in Fig. 2. As shown in Fig. 2, geno-
types in each nuclear family appear to be consistent at the level one
checking, but individual 3 determines the phase (“linkage phase” is
defined as the haplotype of the gamete transmitted from parent to
offspring, see Chapter 1) of individual 6, which forces individual 3
to carry a “2” allele, which also means that a parent of individual 4 is
a carrier of a “3” allele. However, we know the genotypes of the
parents of individual 4, which are 4/2 and 1/2; therefore, he
cannot possibly carry a “3” allele. This type of error can easily be
detected by the genotype-elimination algorithm as a level 2 error).

1.3 The Critical In Subheading 1.2, we described the nuclear-family method and
Genotype Method the genotype-elimination method. Below we discuss two additional
algorithms termed “critical genotype” and “odds ratio method,”
respectively, both implemented in PedCheck.
The genotype of an individual that resolves the issue of incon-
sistency within a pedigree when removed from the data is defined as
a “critical genotype” by O’Connell and Weeks [13].They pointed
out that a “critical genotype” does not always provide a perfect
solution. The critical genotypes may not be independent. For
example, when a parent and offspring are homozygous for different
alleles, both will be critical genotypes, but blanking either resolves
the inconsistency. Therefore, O’connell and Weeks concluded that
“the set of erroneous genotypes is a subset of the critical geno-
types.” The critical-genotype algorithm implemented in PedCheck
enables users to identify the critical genotypes in a particular pedi-
gree by “untyping” one typed individual at a time (meaning call this
individual unknown in the input file), and then applying the
genotype-elimination algorithm to determine if the inconsistency
has been resolved. If one critical genotype is found, this genotype
represents the error, otherwise the set of critical genotypes blanked
may depend on the order of the individuals who were untyped.
Below we give one example to demonstrate how PedCheck
detects errors at different levels. The genotype data were simulated.
PedCheck can be freely downloaded from the following
website: watson.hgen.pitt.edu/register/docs/pedcheck.html. As
documented in the PedCheck menu, using PedCheck one is able
to specify the algorithms at four levels (from basic to comprehen-
sive): level 1 uses the above described nuclear-family algorithm;
Random documents with unrelated
content Scribd suggests to you:
Lifting up the very base 26 of the island E malana i luna i ka ili kai. 15
To float on the surface of the sea. 27 15 Huna e Hina i ka eheu o ka alae,
Hidden by Hina 28 were the wings of the alae. Wahia ka papa ia Laka,
Broken was the table 29 of Laka. A haina i lalo ia Wakea.
Carried far down to Kea, 30 Ai mai ka ia, o ka ulua makele,
The fish seized the bait, the fat, large ulua. 31 O Luaehu, kama a Pimoe, e Kalani e
Luaehu, 32 offspring of Pimoe, O thou great chief! 33 20 O Hulihonua ke kane,
Hulihonua the husband, O Keakahulilani ka wahine;
Keakahulilani the wife; 34 O Laka ke kane, o Kapapaiakele ka
Laka the husband, Kapapaiakele the wife; O Kamooalewa ke kane,
Kamooalewa the husband, O Nanawahine kana wahine; 25
Nanawahine his wife; 25 O Maluakapo ke kane,
Maluakapo the husband, O Lawekeao ka wahine;
Lawekeao the wife; O Kinilauaemano ke kane,
Kinilauaemano the husband, O Upalu ka wahine;
Upalu his wife; O Halo ke kane, o Koniewalu ka wa
Halo the husband, Koniewalu the wife; 30 O Kamanonokalani ke kane,
Kamanonokalani the husband, O Kalanianoho ka wahine;
Kalanianoho the wife; O Kamakaoholani ke kane,
Kamakaoholani the husband, O Kahuaokalani ka wahine;
Kahuaokalani the wife; O Keohokalani ke kane, 35
Keohokalani the husband, 35 O Kaamookalani ka wahine;
Kaamookalani the wife; O Kaleiokalani ke kane,
Kaleiokalani the husband, O Kaopuahihi la ka wahine;
Kaopuahihi the wife; O Kalalii la ke kane,
Kalalii the husband, O Keaomele la ka wahine; 40
Keaomele the wife; 40 O Haule ke kane,
Haule the husband, O Loaa ka wahine;
Loaa the wife; O Nanea ke kane,
Nanea the husband, O Walea ka wahine;
Walea the wife; O Nananuu ke kane, 45
Nananuu the husband, 45 O Lalohana ka wahine; [373]
Lalohana the wife; [372] O Lalokona ke kane,
Lalokona the husband, O Lalohoaniani ka wahine;
Lalohoaniani the wife; O Hanuapoiluna ke kane,
Hanuapoiluna the husband, O Hanuapoilalo ka wahine; 50
Hanuapoilalo the wife; 50 O Pokinikini la ke kane,
Pokinikini the husband, O Polehulehu la ka wahine;
Polehulehu the wife; O Pomanomano la ke kane,
Pomanomano the husband, O Pohakoikoi la ka wahine;
Pohakoikoi the wife; O Kupukupunuu la ke kane, 55
Kupukupunuu the husband, 55 O Kupukupulani ka wahine;
Kupukupulani the wife; O Kamoleokahonua ke kane,
Kamoleokahonua the husband, O Keaaokahonua ka wahine;
Keaaokahonua the wife; O Ohemoku ke kane, O Pinainai ka
Ohemoku the husband, Pinainai the wife; O Mahulu ke kane, 60
Mahulu the husband, 60 O Hiona ka wahine;
Hiona the wife; O Milipomea ke kane,
Milipomea the husband, O Hanahanaiau ka wahine;
Hanahanaiau the wife; O Haokumukapo ke kane,
Haokumukapo the husband, O Hoao no ka wahine; 65
Hoao was the wife; 65 O Lukahakona ke kane,
Lukahakona the husband, O Niau ka wahine;
Niau the wife; O Kahiko ke kane,
Kahiko the husband, O Kupulanakehau ka wahine;
Kapulanakehau the wife; O Wakea la ke kane, 70
Wakea the husband, 70 O Papa ka wahine.
Papa the wife.
Hanau ko ia ka lani he ulahiwa nui.
A chief was conceived and born, a great red fowl. He alii o Pineaikalani, ko kupunakan
A chief was Pineaikalani, thy grandfather, Hanau ka lani he alii;
A chief who begot a chief, Hua mai nei a lehulehu; 75
Bearing innumerable offspring. 35 75 Kowili ka hua na ka lani;
Mixed are the seed of the noble chief, Lele wale mai nei maluna.
Clamoring to be recognized Ka leina a ka lani weliweli.
As being of thy stock, O dread chief. He alii pii aku, koi aku, wehe aku,
A chief ascending, urging on, opening upwards A loaa i ka lani paa ke alii. 80
Until the heaven is reached, 36 where the king is held E Ku e (Kualii), he inoa.
fast. 80 Ina no oe, i ona?
This, O Ku, Kualii is thy name. 37
Dost thou not already stand at its height? 38 O Ku o ke koi makalani!
Kakai ka aha maueleka,
O Ku, thou axe of chiefly edge! 39 Na Ku! kohia kailaomi e Ku! 85
The train of clouds 40 along the horizon doth march Kai Makalii, kai Kaelo,
For Ku, the edge of the sea is drawn 41 down by Ku. 85 Kai ae Kaulua.
The sea of Makalii, the sea of Kaelo, Ka malama hoolau ai a Makalii
The rising sea in Kaulua. O ke poko ai hele, ai iwi na.
The month of Makalii 42 in which the food bears leaf, Ka pokipoki nana i ai ka iwi o Alakap
The worm that eats as it crawls, even to the rib. O ka makua ia o Niele o Lauineniele
The sea-crab 43 that ate the bone of Alakapoki 90 O kanaka o ka wai.
Who was the parent of Niele of Lauineniele, 44 O Ku, ke alii o Kauai.
The people of the water. 45
Ku, the king of Kauai. O Kauai mauna hoahoa,
Kauai with its high 46 mountains. Hohola i lalo o Keolewa. 95
Spread down low is Keolewa, 47 95 Inu mai ana Niihau ma i ke kai-e.
Niihau and the others 48 are drinking the sea. O Kiki ma ka kai Keolewa.
Ah, it is Kiki and his company that are at Keolewa, O Kamakauwahi ma ka kai luna e
Kamakauwahi and his company that are above, O Hawaii.
O Hawaii.
O Hawaii, mauna kiekie. 100
Hawaii of high mountains; 100
Hoho i ka lani Kauiki;
Towering unto heaven is Kauiki. 49
Ilalo ka hono o na moku,
Down at the base 50 of the islands,
I ke kai e hopu ana
Where the sea holds it fast.
O Kauiki.
Kauiki,
O Kauiki i ka mauna 105 [375]
Kauiki the mountain, 105 [374]
I ke opaipai, e kalaina e hopu ana—
Like the sea-gull flapping its wings when about to fall. 51
O Kauai.
Kauai,
Great Kauai inherited from ancestors. 52 O Kauai nui kuapapa,
Noho i ka lulu o Waianae,
Sitting in the calm of Waianae He lae Kaena, 110
Kaena is a point, 53 110 He hala o Kahuku,
Kahuku is hala-wreathed. He kuamauna hono i kehau Kaala,
Covered with dew is the back of Kaala; 54 Noho mai ana Waialua i lalo-e—
There below doth Waialua sit, O Waialua ia.
That is Waialua. O Mokuleia, Kahala ka ipu, 115
Mokuleia with its dish of Kahala; 115 Ka loko ia mano lalawalu,
A fish-pond, like cooked 55 shark,
Hiu lalakea o Kaena,
The tail of the hammer-headed shark is Kaena, Mano hele lalo o Kauai-e—
The shark that travels at the bottom of Kauai, Olalo o Kauai, kuu aina,
At the bottom of Kauai my land; O Kauai—. 120
O Kauai! 120
Ku is sailing to Kauai Ke holo nei Ku i Kauai
To see the worm-eyed oopu of Hanakapiai; E ike i ka oopu makapoko o Hanaka
Ku is returning to Oahu Ke hoi nei Ku i Oahu-e—
To see the transient oopu, 56 I ike i ka oopu kuia, ia
The shameful fish of Kawainui 125 Hilahila o Kawainui 125
Floating near the surface of the water. E lana nei iloko o ka wai.
When the hala is ripe the neck becomes red; 57 A pala ka hala, ula ka ai-e—
’Tis a sign of Ku, He hailona ia no Ku,
He has now landed. O Kauai! Ua pae mai-la-o Kauai!
Great island of lehua is Kauai; 130
Like a moving island in the sea, O Kauai nui moku lehua, 130
Like a moving island toward Tahiti— Moku panee lua iloko o ke kai,
Looking from Tahiti while Wakea 58 has the sun— Moku panee lua ana Tahiti,
Invited, 59 Kona first meets the eye, Halo Tahiti ia Wakea ka la,
Growing up as though from Kumuhonua, 135 Kolohia kau mai ana Kona i ka maka
Rattling the foundations of Hawaii of Kea, 60 Hooulu ilalo o Kumuhonua, 135
Pointing to the early rays 61 of the sun; O nakeke ka papa i Hawaiiakea.
Kona is visible to the eye— O kuhia i ka muo o ka la;
Kona is plainly seen, Kau mai ana Kona i ka maka—
Kohala stretches forth 62 in the distance. 140 Ke kau la Kona,
Tahiti! Whose is Tahiti? Ke moe la Kohala. 140
For Ku indeed. O Tahiti! Ia wai Tahiti?
Tahiti, that island stretched far across the ocean, Ia Ku no.
Land where Olopana 63 once dwelt.
Within is the land, without is the sun; 145 O Tahiti moku kai a loa,
Approaching that land the sun hangs low. 64 Aina a Olopana i noho ai.
Perchance you have seen it? Iloko ka moku, iwaho ka la; 145
I have seen it— O ke aloalo o ka la ka moku ke hiki
I have indeed seen Tahiti, 65 Ane ua ike oe?
Tahiti where the language is strange. 150 Ua ike.
To Tahiti belong the people who ascend
Ua ike hoi wau ia Tahiti.
To the backbone 66 of heaven,
He moku leo pahaohao wale Tahiti.
And while above they tread
No Tahiti kanaka i pii a luna
And look down below.
A ka iwikuamoo o ka lani;
There are none like us in Tahiti. 155
A luna, keehi iho,
Tahiti has but one kind of people, the haole. 67
Nana iho ia lalo.
They are like unto gods;
Aole o Tahiti kanaka; 155
I am like a man. Hookahi o Tahiti kanaka, he haole.
A man indeed, Me ia la he akua,
Wandering about, and the only one who got there. 68 160 Me a’u la he kanaka;
Kukahi is past, Kulua comes next; [376] He kanaka no.
Kukahi the night, Kulua the next day. Pai kau, a ke kanaka hookahi ia e h
The food is partly eaten, Hala aku la o Kukahi la o Kulua, [377
Eaten as by birds little by little. 69 O Kukahi ka po, o Kulua ke ao,
Listen now, bird of victory! 165 O hakihana ka ai;
Victory! Through whom? Kanikani ai a Manua.
Through Ku indeed. Hoolono mai Manuolanakila, 165
The rain was there, the sun was there, Malie ia wai lanakila?
The early rising star 70 was there, a king. Ia Ku no.
Kaulakahi the sun, 170
Kapukahi the day, Ilaila ka ua, ilaila ka la;
Puna, hooilo, 71 Hana, lanakila; Ilaila ka hoku Hikimakahano he alii.
Hooilo, Pele 72 is angry. O Kaulakahi ka la, 170
The wind! Who hath the wind? Kaupukahi ka la,
It is held by Ku indeed. 175 O Puna, o hooilo, o Hana, o lanakila
Blown is the wind of Laamaomao, 73 O Hooilo, ua ino Pele.
Like the soft koolauwahine 74 breeze below. O ka makani! Ia wai ka makani?
Kauai have I seen, Ia Ku no. 175
The northwest wind of Wawaenohu,
Puhia ka makani a Laamaomao,
The north wind of Niihau, 180
Ke ahe koolauwahine ka makani o la
The south wind is the strong wind,
O Kauai ka’u i ike,
The aoa 75 the tempestuous wind,
O ke kiu ko Wawaenohu,
The wind scattering kukui blossoms on the flood, 76
O ka hoolua ko Niihau, 180
Carried to awaken Lonomoku, 77
O ke kona ka makani ikaika,
Blowing there below at Hanae, 185
O ka aoa ka makani ino,
For of such is the koolauwahine of lower Kauai
Ka makani halihali wai pua kukui,
While it tarries at Wailua. 78
I lawe ia la e Lonomuku,
The star! Whose is the star?
Pa ilalo o Hanae 185
For Ku indeed.
Oia koolauwahine o lalo o Kauai
The rain is at Puanalua. 79 190
E pa ka i Wailua la la.
Pierced by the three stars 80 of Orion
O ka hoku! ia wai ka hoku?
Are the clouds as they drift 81 on.
Ia Ku no.
The stars are peeping out two by two,
The south wind and rain. Iluna ka ua o Puanalua 190
Pierced by the three stars of Orion 195 Ku i ke kao Maaiku hoolewa
Are the clouds as they drift on. Ka wae o ke kaina,
The rain! Whose is the rain? Oiliili lupea na hoku mahana elua.
For Ku indeed. He ua kona me ka makani,
Slantingly 82 falls the rain at Kunaloa, Ku i ke kao Maaiku hoolewa 195
Dropping singly 83 on the skin; 200 Ka wae o ke kaina
Heavily falls the rain at Kananaola; 84 O ka ua, ia wai ka ua?
Slippery is Mahiki, the traveler falls. Ia Ku no.
For Hina hath sprinkled the rain,
Which shook the bottom of Maheleana, I moea ka ua i Kunaloa,
The mist of the rain 85 was at Kahalahala, 205 I pakakahi ka ua i ka ili, 200
The small rain 86 Iliki ka ua i Kananaola,
That falls amongst the lehua. Pahee Mahiki ke ka la,
The sun! Whose is the sun? Ua lu ia ka ua e Hina,
For Ku indeed. Haalulu ai lalo o Maheleana.
The sun comes forth at Kauiki; 87 210 O ka punohu o ka ua kai Kahalahala
Scorching 88 is the sun at Upilialoula. O ka pokii o ka ua,
The children are piping forth 89 E ua ka i ka lehua la,
The sun is sinking, sinking fast, O ka la, ia wai ka la?
The sun 90 in the flower-nets of Hilo. Ia Ku no.
The back of the sun is turned up, 215
The face of the sun is turned below, I puka ka la ma Kauiki, 210
The shade of the sun is turned inward, Hawewe ka la i ka Upilialoula,
The light of the sun is traveling over the land Ke kohokoho la kamalii,
And sinks beyond Lehua. 91 Ke na’u na’u la ka la,
The sea! Whose is the sea? 220 Ka la kieke pua o Hilo,
For Ku indeed. O ke kua o ka la kai hulihia iluna, 21
The vastness of the sea is from Tahiti, 92 [378] O ke aloalo o ka la kai lawea ilalo,
Calm 93 is the sea by the land, O ka malu o ka la kai kaa iloko,
Taken up is the sea in the hand. O ke aka o ka la ke hele nei maluna
Dressed 94 is the hair with the sea, 225 A kau aku i Lehua la la.
Red is the hair with very salt 95 sea, O ke kai! ia wai ke kai? 220
Softened is the hair in the great sea, 96 Ia Ku no.
Brown is the hair 97 with the foamy sea. I nui mai kai i Tahiti, [379]
Rich is the soup 98 of the hog, I miha kai i ka aina,
Fat is the soup of the dog, 230 I lawea kai i ka lima,
Dainty the soup of the fowl, I hiki ke oho i ke kai, 225
Fragrant the soup of the mullet, I ehu ke oho i ke kai liu,
Strong the soup of the palani. 99 I pala ke oho i ke kai loa,
The sea for surf-riding is at Kahaloa, 100 I lelo ke oho i ke kai kea.
The sea for casting the net is at Kalia, 235 He kai kuhikuhinia ko ka puaa,
The sea for going naked 101 is at Mamala, He kai lihaliha ko ka ilio, 230
The sea for swimming is at Kapuuone, He kai okukuli ko ka moa,
The sea for surf-riding sideways 102 is at Makaiwa, He kai ala ko ka anae.
The sea for kicking up mullet 103 is at Keehi, He kai hauna ko ka palani,
The sea for small crabs 104 is at Leleiwi, 240 He kai heenalu ko Kahaloa,
The sea of many harbors 105 is at Puuloa, He kai hului ko Kalia. 235
A sea that blows up nehu and lala, 106 He kai hele kohana ko Mamala,
Is the sea of Ewa so calm; He kai au ko Kapueone,
The great Ewa [lands] of Laakona. 107 He kai kaha-nalu ko Makaiwa,
Ku 108 holds up the rain in the heaven. 245 He kai ka anae ko Keehi,
The mottled 109 sea of Heeia, He kai elemihi ko Leleiwi, 240
The sea for spearing hee is at Kapapa, He kai awalau ko Puuloa,
The sea that calls 110 is at Kualoa, He kai puhi nehu, puhi lala,
The sea that wears away is at Kaaawa, Ke kai o Ewa e noho i ka lai nei,
The wild sea 111 is at Kahana. 250 Na Ewa nui a Laakona,
Paao 112 left open the sea, Ku i ke alai ka ua i ka lani, 245
Paao is seen in the waterfall, Kai apukapuka Heeia,
The great one is known from below Kona, He kai ohee ko Kapapa,
He is the great 113 one from below Kona. 114 He kai oha i ke Kualoa,
A handle, an axe, the cord, the cover; 115 255 He kai aai ko Kaaawa,
He kai ahiu ko Kahana. 250
Take it, bind it, wind it round, 116 I wehe kai ia Paao,
And cut down the foundations of Tahiti. Ikea Paao i ka waihi,
While it rains at Hilo Ikea ka hiwa mai lalo Kona,
The rain-clouds at Maheleana are in conflict 117 with one O ka Hiwa ia, mai lalo Kona,
another, He a’u he koi, he aha he pale, 255
Else it will rain on the people. 260 E kii e hoa e lanalana,
For there the rain is till it ceases; E kua i kumu o Tahiti-e,
A long day 118 in the wind, O ua mai Hilo.
Cramped is 119 the traveler by the rain. Ke kuee nei na opua ua o Mahelean
Mahiki causes struggling 120 with its muddy hills; O ua mai kanakea, 260
There Mahiki causes him to fall. 265 Ilaila ka ua a malie,
Lola loa ai ka makani,
Haiki ka make a ka ua,
Hakookoo ana Mahiki i ka puukolea,
Aia Mahiki ke ka la. 265

While Kapaahulani was chanting this section of the mele I loko o keia wa a Kapaahulani e heluhelu la i
Kualii’s chief officer remarked to the king: “Why are we ka pukaua o Kualii iaia: “E apa ana kakou i kei
delaying this battle? Why not let us fight out this small noa he hoouka ae ka kakou i keia wahi kaua u
battle?” Kamakaaulani, the brother of Kapaahulani, then olelo aku o Kamakaaulani, ke kaikaina o Kapaa
spoke to the chief officer: “Let us perhaps delay the “E kali paha ka hoouka o ke kaua, ke loaa mai
battle. I hear the king’s name is being mentioned; ke alii; nolaila e pono kakau ke kali aku; ina i h
[380]therefore it is but proper that we wait. If he can Kualii mau kupuna mai ka po mai a hiki ia iane
rightly name Kualii’s ancestors from the beginning right hookuu aku, aole e kaua.” A e like me ka olelo
down to him (Kualii), then we should let them go, and pela no o Kualii i hoolohe ai. Aka, o Kapaahula
there will be no fighting.” Kualii, by his silence, showed kana hana ana i keia mele, ka mea i kapaia he
that he would abide by the words spoken by
Kamakaaulani. In the meantime Kapaahulani continued
on with his chanting of the mele, called a prayer.

Opuukahonua, 121 Lolomu, Mihi, O Opuukahonua, o Lolomu, o Mihi,

Lana the wife. O Lana ka wahine.
Wakea lived and took Papa, Noho Wakea noho ia Papa,
Lived with Kanananuukumamao. Noho ia Kanananuukumamao,
Naupaka 122 was then born, which stands by the Hanau ka naupaka, ku i ke kahakai,
seashore. 270 O Ohikimakaloa ka wahine,
Ohikimakaloa the wife, O Hoopio, o Hulumaniani,
Hoopio, Hulumaniani, Ku i ka ena, anaia ilalo.
Overcome by jealousy 123 O Mehepalaoa,
He was struck down. O Naholo, 275
Mehepalaoa, 124 Mehe kai olohia a Manu,
Naholo, 275 Oia alakai honua Ku.
Like the broad calm 125 sea of Manu, O Lanipipili, O Lanioaka,
The one who led 126 Ku to earth. O Lanikahuliomealani,
The closed heaven, 127 the open heaven, O Lono, o Hekilikaaka, 280
The heaven that turns, the heaven that breaks up. 128 O Nakoloailani,
Lono the rolling thunder, 129 280 O Kailolimoana, O Waia,
The heaven that rumbles 130 O Hikapoloa,
The disturbed sea. 131 Waia. O Kapoimuliwaa,
The long-delayed night, 132 O Kane, 285
The last of the nights, O Ahulukaaala,
Kane 133 285 O Kaneikamakaukau,
And Ahulukaaala; Alua anahulu wau ia oe e Ku-e;
Kane who is ever ready. 134 E Kualii.
Twice ten days 135 am I with you, O Ku, Eia ke paia ai o Kapaau, 290
O Kualii. He kanaka o Wawa kai kapua.
Here is the attractive hook 136 of Kapaau; 290 Kea pua ko o Hawi,
The men of Wawa 137 are at Kapua, Eia ke puhi kukui ai o Kukuipahu,
Playing with the sugar-cane arrows of Hawi. Ka wahine wahaula ke ai i ka ina o M
Here is the torch of Kukuipahu, 138 Eia ke kanaka pii pali 295
The woman whose mouth is made red by eating the Haka ulili o Nanualolo,
sea-urchin 139 of Makakuku. Keiki kia manu-e—
Here is the cliff-climber, 295 Kau kia kia manu o Lehua. [383]
Of the ladder of Nanualolo, 140 O Kuku, o Aa, o Naio,
The bird-catching youngster O Haulanuiakea ke koi, o Hinaimana
Raising his bird-catching 141 pole at Lehua. [382] O Paepae, o Manau ka wahine,
Kuku, 142 Aa, Naio, Hanau ka naenae noho kuamauna
Haulanuiakea, 143 the axe of Hinaimanau, 300 Ka hinihini kani kuaola,
Paepae, Manau his wife, Hakina iho i ka wae mua,
They brought forth Kanaenae 144 that dwells on the O Molokai ua naha ke ’na, 305
mountain, Haalele aku Kanaloapuna,
The hinihini 145 that sings on the high mountain. Kanaloa a Waia,
Broken on the front 146 seat, O Make holo uka,
That is Molokai torn asunder, 147 305 O Make holo kai.
Deserted by Kanaloapuna— Hoonalulu ana Luukia, 310
Kanaloa of Waia. Hoopailua i ka iloli,
There is death if you run toward the mountains; I ke kauhua o ke kamaiki
There is death if you run toward the sea. Hanau ka ieie hihi i ka nahele,
Luukia is suffering headache, 310 O Makaaulii kana wahine.
Made sick by the unpleasant sensation of pregnancy Hanau ka lupua me ka lau lama, 315
Conceiving the child. Ku i ke opu o Lono.
The ieie 148 is conceived that creeps in the forest, O Kapolei ka wahine,
Makaaulii was his wife O Ku kaikaina i hope ka lanalana,
Which brought forth the lupua and laulama 315 O Kukonaihoae, o Ku,
Like unto the bushy stock of Lono, O ke kai mahuehue, 320
Kapolei was the wife. Mehe kai e haa aku ana Ku.
Kukaikaina behind the spider, Eia ka wahine peeki
Of Kukonaihoae, Ku. Uhi lepo o Keaau,
The rising sea 320 Ka umeke hoowali ’na lepo,
Like unto a dancing sea is Ku. Mehe hako la ke ala, 325
Here is the woman that hides, Eia na huakai hele
Covered by the dust of Keaau, 149 Alanui ka kanaka.
The calabash of kneaded earth. Wali ai ka lepo o Mahiki,
Like unto the leaf of the sugar-cane is the path. 150 325 I ka paala a ka wawae.
Here is the company of travelers. O Kapapaiakea, o ka nalu o ka inain
The slippery road that makes men fall, O Kauhihi i kana wahine,
Which softened the dirt of Mahiki, 151 Hanau koa waa ku i ka nenelu,
Being trodden down by the foot. Kalaia ka ipu i ke kai aleale,
Kapapaiakea the wave of wrath, 152 330 Kalaia o Hinakapeau,
Kauhihii was his wife Loaa mai o Ukinohunohu la, 335
Which brought forth the koa for canoes 153 growing in Ukinaopiopio, o Moakueanana,
the rich soil. O Kalei, o Keelekoha,
Loosened was the vessel in the rippling sea, O ke ’kua maka holo lalo,
Loosened was Hainakapeau. 154 O ke kau i luna Kahualewa,
Then was seen Ukinohunohu, 155 335 Ako lipoa o Kanamuakea, 340
Ukinaopiopio, 156 Moakueanana, O ke kai akea
Kalei, Keelekoha, 157 O ka moana akea.
The god with the downcast eyes. O Hulukeeaea
The turned-up [eyes] of Kahualewa, O Hauii, o Hauee,
Gathering the seaweeds of Kanamuakea, 340 O Hauii nui naholoholo, 345
The broad sea, O Hauii kai apo kahi,
The broad ocean Kai humea mai ko malo e Ku.
Of Hulukeeaea. 158 No Ku ka malo i ke kaua haa oe.
The cold dew, the shivering dew, Oia i luia, ka umu mehe awai la.
The great dew that makes one quiver, 345 Eia ka uhuki hulu manu, 350
The cold dew that comes 159 over the sea, Kau pua o Haili,
The sea where you put on the loin cloth, 160 Ku. Na keiki kiai pua,
Ku puts on his loin cloth for war, and you tremble. Ka lahui pua o lalo.
Scattered is the oven, 161 like the rushing of a stream. Eia ka wahine ako pua, [385]
This is the plucker of feathers, 350 Kui pua, lei pua, kahiko pua o Paiah
The bird-catcher of Haili, 162 Ke uhai mai nei i ke ’kua,
The boys that watch the flowers, A pau mehameha Apua.
The bunch of flowers that is below. Kauia ka makani, hiamoe-la-e—
This is the woman that picks the flowers, [384] Moe ua makani, hiamoe la la-e
Stringing flowers, wearing flowers, dressed in the I ka papa o Kukalaula. 360
flowers of Paiahaa, 163 355 O Uliuli, o Maihea,
Who comes chasing after the ghosts. 164 O Kahakapolani ka wahine,
It is gone, all is deserted like Apua, O Kaukeano, o Mehameha,
The wind of the sleep that wakes not 165 has passed over, O po ka lani i ka ino;
The wind that sleeps, sleeps on, He ino ka lani ke wawa nei ka honu
On the broad expanse of Kukalaula, 166 360 I ka inaina o kalani.
Uliuli, 167 Maihea, 168 Hoonaku, hookaahea, hoowiliwili,
Kahakapolani the wife, Hoonahu, hoomamae,
The sacred, the lonely place. 169 Hookokohi ana iloko o Hinaiaeleele.
Dark is the heaven with storm, Hanau ka maua Ku i ka nahele, 370
The heaven is stormy, for the earth is rumbling, 170 365 Hanau ka ouou kani kuaola,
Caused by the water that precedes 171 the birth of the Puka ke kamahele
king, Ku i ke alo o ka hakoko.
Travailing, fainting, struggling, He pukaua na ke alii, he kaua.
Suffering pangs, feeling the pressure, He wai kaua o Ku no ke alii, 375
Bringing forth in the month of Hinaiaeleele, He kaua na Ku,
Our Ku is brought forth in the forest, 370 E uhau ana iluna o Kawaluna.
Brought forth the ouou 172 singing on the mountain ridge. Ihea, ihea la ke kahua,
The child is brought forth; Paio ai o ke koa-a?
It stands in the presence of the travailing mother, I ka i kahua i Kalena, 380
A chief warrior for the king, an army, I Manini, i Hanini
An army of hosts, for Ku is indeed king. 375 I ninia i ka wai akua,
A battle for Ku, I ko hana i Malamanui
Beating his enemy on the heights 173 of Kawaluna. Ka luna o Kapapa, i Paupauwela,
Where? Where is the battle field I ka hilinai i ke kalele, 385
Where the warrior is to fight? Ka hala o Halahalanui maauea,
On the field of Kalena, 380 Ke kula ohia ke Pule-e,
At Manini, at Hanini, Ke ’kua o Lono o Makalii
Where was poured the water of the god Ka lala aalao Ukulonoku,
By your work at Malamanui; No Kona paha, no Lihue. 390
On the heights of Kapapa, at Paupauwela, No ka la i Maunauna,
Where they lean and rest; 385 No ka wai i Paupauwela,
At the hala trees of indolent Halahalanui, I ulu haa lilo i Nepee,
At the ohia grove of Pule-e, 174 A ka hauna o Aui.
The god of Lono, of Makalii, Kikomo kahuna i kakua laau, 395
The fragrant branch 175 of the Ukulonoku, Komo Ku i kona ahuula,
Mayhap from Kona, 176 from Lihue, 390 Ka wela o ka ua i ka lani,
For the day at Maunauna, 177 Ka la i Kauakahihale, [387]
For the water at Paupauwela. Ula ka lau o ka mamane,
Growing low at Nepee, Ke koaie o Kauai; 400
At the slaughter 178 of Aui, He pili kai ihi ia e Ku,
Where the priests joined in the battle; 395 Ka aloalo o Kamaile,
Ku is arrayed in his feather cloak; 179 Ka nalu kakala o Maihiwa,
The sun-lighted 180 rain in the heavens; Pania ka wai i Halapo,
The sun at Kauakahihale. 181 Ka naha ilalo o Eleu. 405
[386] Huki kaua a moa i ka lani,
Red is the leaf of the mamane, 182 Me he hee nui no kuahiwi;
The koaie 183 of Kauai; 400 Ka hee na o Hilo ia Puna,
The sea grass has been stripped 184 by Ku— Aia ma Hilo Peahi;
The waving [grass] of Kamaile; Ula ka wai i Paupauwela, 410
The towering surf 185 of Maihiwa, Ke kilau o Malamanui,
Which dammed 186 up the waters of Halapo. Ka moo kilau i Kapapa.
The breaking up is below at Eleu, 405 Kui ka lono ia Haalilo,
The rain is drawn away to the sky, Haua aku la ko kaina;
Like a full retreat from the mountain; Hahaki Haalilo i ka manawa; 415
It must be the defeat 187 of Hilo by Puna, I kai muku kahuna ia Ku;
There at Hilo is Peahi. I la ka mawa ia Ku,
Red is the water of Paupauwela, 410 I keiki a Haalilo.
From the slain at Malamanui, Eia Malanaihaehae,
The slain on the ridge at Kapapa. Kama a Niheu kalohe, 420
The tidings 188 reached Haalilo Ke pani wai o Kekuna,
Your younger brother 189 is beaten. He mee nei no ke kanaka,
Haalilo is sore at heart, 415 Ke pu nei i ka aahu,
For Ku has left but few of the priests; 190 Ke lapa nei i ka lauu,
They are beaten 191 by Ku, Ka laulau o ka palau, 425
The children of Haalilo. Eia Haalilo-e!—
Here is Malanaihaehae, 192 O Ku no ke alii.
Offspring of mischief-making Niheu, 193 420 Aloha kukui peahi i na leo paoa;
The dammer of the waters of Kekuuna. Ua oa ka maka o ka ilima make,
A prodigy 194 among the people. Nonu i ka malama o Makalii; 430
He is girding 195 on his robe, Ia Makalii la pua ke koolau,
He is whirling 196 his weapon [in the air], Pau i ke hau o Maemae.
The war club is caught in his 197 robe. 425 He mae wale ka leo o ke kai o lalo,
Here is Haalilo, 198 Hoolono uwahi o Malamanui,
Ku is indeed king. Ia ai ku i ka uwala, 435
Dear are the kukui trees wafting 199 the message of Kauwewe kupukupu ala o Lihue.
Paoa, Kupu mai nei ka manawaino e Ku-e
The tender shoots of the ilima are dead, Hanau mai a me Kalani wale la;
Withered 200 in the month of Makalii. 430 O Ku no ke alii.
It is in Makalii that the koolau 201 blooms,
Wet with the dew of Maemae. He pu hinalo no Ku i Kamakoa, 440
Fast fades the voice of the sea 202 there below; Oi lele Ku i ka pali,
Malamanui listens for its voice, Mai pau Ku i ke ahi,
Where Ku ate the potato 435 O keaha la kau hala e Ku?
Covered in cooking with the fragrant kupukupu 203 of O ke kua aku i ka laau,
Lihue. O ka luukia ana o ka pau, 445
Ugly thoughts are breeding, O Ku, O ke ahina ana o ke oa,
It is brought forth and towers to heaven; O ko Ku ia o kona hoa haele
For Ku is indeed king. Kaua i ka la.
Aai ai Ku i ka unahi pohaku,
The voice of Ku was like the trumpet of hala blossom at Ola Ku i ka ipu o Lono 450
Kamakoa, 440 I ka ipu a Kupaka,
While Ku jumped down the cliff. O Ku no ke alii. [389]
Well nigh perished was Ku by fire. O Kailua makani anea oneanea,
What is thy failing, O Ku? Makani aku a Hema,
Is it the cutting of the tree? He mama wale ka leo ke uwalo mai
Is it the binding 204 of thy robe? 445 E o ianei o Kalahuimakani,
Is it the thrusting of thy spear? 205 E ku mai oe i ka hea i ka uwalo,
The spear, the companion of Ku Mai hookuli mai oe;
In the sun, in the rain? O ke kama hanau
Where Ku did eat 206 the scale of the rock. O ka leo kai lele aku la i waho, 460
Ku was saved by the vessel of Lono, 450 Kai no i waho ka paio,
The vessel of Kupaka, Pale aku la ilaila;
For Ku is indeed king. Hoio mai ika hale liliia,
[388] Mehe leo la ko ka aha,
Kailua with its dry waste wind, 207 Ke kaunuia la ka moena, 465
The wind sent by Hema, Ke kapa me ka aahu,
Where the voice sounds weak 208 in its pleadings; 455 Ke hea wale la i ka uluna-e
The voice which is still ringing till it reaches Aole ia he kanaka.
Kalahuimakani. O maua no na kanaka.
Stand forth at the call, at the pleading;
Turn not a deaf ear.
By the birth of offspring, 209
The voice hath indeed gone forth. 460
We had thought the battle was to be without
Where we would indeed contend it.
But you return to the house 210 with jealousy,
As though the battens 211 could respond;
You show anger 212 to the mats, 465
The bed covering, and the robes;
And call to the pillows in vain.
That is not [the actions of] a man;
We are indeed the men.

When Kapaahulani reached that portion in his chant I keia manawa, a Kapaahulani i hea ai: “E ku m
where it says, “Stand forth at the call, at the pleading; ualo, mai hookuli mai oe.” Ma keia kaukau ana
turn not a deaf ear,” Kualii was raised up while still in the la o Kualii me ka paa i ke koko; na Kualii i ku m
netting. 213 As he was being held up, the soldiers of the me na koa o na aoao elua o ko Kualii puali no;
two wings of his army also stood up. When the warriors ke kaua o Koolauloa mai, aole i kana mai a ka
from Koolauloa looked about them and saw that Kualii Kualii. Ia manawa, iniki aku, iniki mai ka puali
had a very large army, they nudged back and forth and me ka olelo iho: “Ike wale ke kahuna; eia ka u
remarked to one another: “The priest 214 is indeed wise; kaua.” Alaila ia wa, ke nalu la na pukaua o ko
here we are indeed surrounded by the enemy.” At this i ka pilikia.
same time the chief warriors of the Koolauloa army were
thinking of the trouble they had gotten into.

While Kualii was being held up in the koko, standing I kela manawa a Kualii e amo ia la iloko o ke k
amidst his warriors, Kapaahulani still kept on chanting hooki o Kapaahulani i kona kahea ana i ka ino
the mele of Kualii, never once stopping to rest, from the o kana kahea ana mai no mai kinohi mai, o ka
time he began it until both armies met. When me ka hoomaha ole a hiki i ka hui ana o na ka
Kapaahulani saw Kualii he endeavored to find some one
with whom he could compare Kualii, but try as he would Ia Kualii i kela manawa, hea mai la o Kapaahu
he was unable to find any one worthy of comparison hoohalike ana ia Kualii i mea e like ai, aole nae
except Keaweikekahialiiokamoku, the king of Hawaii. Aka, o ka mea i like ai o Kualii ma ka hoohalik
The reason of this was because Kualii was compared to Keaweikekahialiiokamoku ke alii o Hawaii. Nok
a god. 215 When Kualii walked from Kahaluu to Waialua akua o Kualii. Ina e hele o Kualii mai Kahaluu
by way of the sea, the sea within the reef would dry up. maloko o ke kai, alaila maloo ke kohola.

Keaweikekahialiiokamoku, 216 on the other hand, when A o Keaweikekahialiiokamoku hoi, ina e holo o
out fishing with his men and ran out of water would turn kanaka i ka lawaia, a ina i makewai na hoewaa
salt water into fresh. When his paddlers became thirsty Keawe: “E! makewai makou.” Alaila ukuhi ae la
they would tell Keawe: “Say, we are thirsty.” The iloko o ka waa, alaila hoi ala mai la hoi o Keaw
paddlers would then dip up the sea and pour it into the ka pola o na waa a noho i ka noho-ana, hoole
canoe, and Keawe would get up from the covered seat o ke kaina liu, alaila, ua manalo koke ke kai, a
between the two canoes and sit in one of the seats; na hoewaa; pela i lilo ai o Kualii i mea like me
then he would stretch out his legs in the sea water that Keaweikekahialiiokamoku.
had been poured into the canoe, and the sea water
would lose its saltness and become fresh, and the
paddlers would then use it for drinking purposes. This
was the reason why Kualii was compared to
Keaweikekahialiiokamoku.

Following is the mele to that effect: [390] Eia malalo iho ke mele o ia ano: [391]

Thou canst not be compared 217 to the twisted hala, 470 Aole i like i ka hala wili, 470
The crooked 218 naio tree, Ke naio laau kekee,
Nor the thick garland of the motherless ahihi, 219 Ka auka a hihi ku makua ole,
Nor the deep pool trod 220 by the bathers, Ke kawakawa i keekeehia,
Nor the hinahina 221 in the wind, Ka hinahina i ka makani
Bending to lean and to fall. 475 Hele ana e hio, e hina-la— 475
Not like these art thou, Ku. Aole i like Ku.
Perhaps thou art like the ohia, 222 Ua like paha ka ohia,
The lehua in the very ninth 223 recess of the forest, Ka lehua i ka wao eiwa,
The tree standing alone 224 in the jungle. Ka laau hao wale ku i nahelehele,
Not like these art thou, Ku. 480 Aole i like Ku. 480
Not like the ekaha 225— Aole i like i ka ekaha,
The ekaha placed under the mat I ka ekaha ku i ka moena,
Like the kiele in fragrance, Me ke Kiele, me ke ala,
Like the changing leaf of the olapa, 226 Me ka olapa lau kahuli,
Like the flower of the fragrant grass, 485 Me ka pua mauu kuku, 485
Falling now hither now thither so easy. Hina wale, hina wale-la—
Not like these art thou, Ku. Aole i like Ku.
[Nor] the rain that brings the land breeze, 227 Ia ua hoohali kehau,
Like a vessel of water poured out. Mehe ipu wai ninia la,
Nor to the mountain breeze of Kumomoku, 490 Na hau o Kumomoku; 490
[The] land breeze coming round to Leleiwi. 228 Kekee na hau o Leleiwi,
Truly, have you not known? Oi ole ka oe i ike
The mountain breezes, that double up your back, I ka hau kuapuu.
[That make you] sit crooked and cramped at Kaimohala, Kekee noho kee, o Kaimohola,
The Kanehili at Kaupea? 495 O Kanehili i Kaupea-la 495
Not like these art thou, Ku. Aole i like Ku.
Not like the lipoa 229 (sea weed), Aole i like i ka lipoa,
The nanue, food of the fishes, Ka nanue ai a ka ia,
The lipahapaha of Waimea, Ka lipahapaha o Waimea,
The moss that hangs on the wood, 500 Ka limu kau i ka laau, 500
The red crab 230 on the top of Kaala. Ka elemihi ula i ka luna o Kaala-la
Not like unto these art thou, Ku. Aole i like Ku.
Not like the kukui, Aole i like i kukui,
The rough-barked kukui, 231 I kukui ili puupuu,
The bark cracked up by the sun. 505 Ili nakaka i ka la, 505
Like to a man who drinks awa, 232 Mehe kanaka inu i ka awa la,
Are the roughness of the kukui trees of Lihue. Ka mahuna o kukui o Lihue la,
Not like these art thou, Ku. Aole i like Ku.
Not like the aalii, 233 Aole i like i ke aalii,
The fragrant poholua tree, 510 Ka poholua laau ala, 510
Nor the maile 234 that grows on the heights 235 of Maoi, Ka maile hoe hoi i Maoi,
Nor the kaluhea 236 of Kawiwi. Ke kaluhea o Kawiwi la,
Not like these art thou, Ku. Aole i like Ku.
Not like the kokio 237 [tree],
With its branches of solitary flowers, 515 Aole i like i ke kokio,
Dropping them wilted to the ground. 238 I ka hahaka pua maoia, 515
Not like these art thou, Ku. Ke kahuli pua i ka paka-la.
Not like the kawau 239 Aole i like Ku.
Is the kalia 240 standing in the open.
Aole i like i ke kawau,
The division recognized a man, 520
I ke kalia ku ma ka waha,
An island, a district, a land. 241
Ai mai ka mahele he kanaka, 520
Not like these art thou, Ku.
He moku, he au, he aina la,
Nor art thou like the porpoise [392]
Aole i like Ku.
With his snout that cuts the sea,
His body in the water where dwells the shark. 525 Aole i like i ka naia, [393]
Not like these art thou, Ku. I kona ihu i kihe i ke kai,
Nor art thou like the kokio, Kona kino i kai; o ka mano la 525
The hawane, nor [the] lehua-flower eater,
Aole i like Ku.
The oo, 242 bird of Kaiona. Aole i like i ke kokio
Not like these art thou, Ku. 530 I ka hawane, ai pua lehua
Not like the pig 243 Ka oo, manu i Kaiona-la
Discerning the progeny of the god; Aole i like Ku. 530
[Or] The breadfruit planted by Kahai. 244
Aole i like i ka puaa,
Truly, have you not known I ka weke lao a ke akua,
The woman with the dyed 245 garment, 535 Ka ulu kanu a Kahai;
On the top of Puuokapolei? Oi ole ka oe i ike,
Not like these art thou, Ku. Ka wahine pau mao 535
Not like the wiliwili 246
I ka luna o Puuokapolei-la?
Whose seeds are made into bracelets, Aole i like Ku.
Whose trunk is gliding; 540 Aole i like i ka wiliwili,
Whose body is down mid the rollers, surf-riding. Kona hua i kupee ia
Not like these art thou, Ku. Ka oiwi ona i hee-a, 540
Not like the beating of the wind.
Kona kino i kai o ka nalu la, heena
Moaning through the mountains, Aole i like Ku.
Bracing up the house of Koolau, 545 Aole i like i ka pa a ka makani,
Fastening it lest it fall by the wind; E nu ana i ke kuahiwi,
The fish-pole of the fisherman, Kakoo ana ka hale o Koolau, 545
The pearl fish-hook of Manaiakalani. 247
Lawalawa ana a hina i ka makani,
Not like these art thou, Ku. Ka mokoi hoolou a ka lawaia,
Not like the mamaki, 248 550 Ka pa o Manaiakalani-la,
With its long, tangling shoots in the forest; Aole i like Ku.
The discerning eyes amid the shrubs, Aole i like i ka mamaki, 550
The small piercing eye, blanched eye;
I ka hialoa maka o ka nahele,
Blanched in the presence of Kuehu, Ka maka kohikohi laalaau;
Along the sacred road, along the long road, 555 Ke a maka ulii, maka ehu,
Along the highway traveled by him. I ehu i ke alo o Kuehu,
Not like these art thou, Ku. I ke ala ihi, i ke alaloa, 555
Not like the ti leaf 249—
I ke alaloa e heleia la-la,
The yellow ti leaf of Nuuanu, 250 Aole i like Ku.
Softened by the rain and wind till it falls; 560 Aole i like i na laki,
The yellow ti leaf on the heights of Waahila. 251 I ka laki pala o Nuuanu,
Not like to these art thou, Ku. I heheia e ka ua e ka makani a he
Not like the Waahila [wind],
Ka laki pala i ka luna i Waahila-la.
The cold wind of Kahaloa, Aole i like Ku.
Scattering the blossoms of the kou, 252 565 Aole i like i ka Waahila.
Stringing them into garlands and carried Ia makani anu o Kahaloa,
To wreath the sea of Kapua. E lu ana i ka pua kou, 565
Not like these art thou, Ku.
E kui ana a paa ia,
Not like the red kamani, E lei ana i ke kai o Kapua-la,
The bright catcher of birds, 570 Aole i like Ku.
Nor the bundle of garlands 253 of hala, Aole i like i ka maniula,
[Or] like the blossoms of the kaa [vine], Ma ke kia ula o na manu la, 570
Whose leaves make music for Ku,
Me ka pa lei o ka hala-la,
For Ku indeed is chief. Me ka pua o ke kaa,
Not like these art thou, Ku. 575 Lau kani o Ku-la
Not like the makole, 254 O Ku no ke alii,
That moisture-laden plant Aole i like Ku. 575
With its creepers hanging over the cliff. Aole i like i ka makole,
Not like these art thou, Ku. Ia laau kewai nui,
Yet thou hast an equal. 580 E hihi ana e ka lihilihi-la
There is a person to whom thou art like: Aole i like Ku.
Keaweikekahialiiokamoku; [394] Ua like; 580
To Keawe, lord of Hawaii. Aia ha kou hoa e like ai,
There is saltness in the sea water, O Keaweikekahialiiokamoku, [395]
There is freshness in the water, 585 O Keawe, Haku o Hawaii-la.
There is heat to the sun, He awaawa hoi ko ke kai,
There is warmth to my skin, He mananalo hoi ka wai, 585
From my husband, Nininini, [is] the water He welawela hoi ko ka la,
Pulele. Are you alike? He mahana hoi ko kuu ili
Not like to these art thou, Ku. 590 Ko kuu kane o Nininini ke wai
Not like this chief O Pulele la. Ua like?
Is there any comparison. Aole i like Ku. 590
He is a man (human), Aole i like nei lani,
Ku is a god. I ka hoohalikelike wale mai;
Ku is a messenger sent from heaven, 595 He kanaka ia,
Ku is a haole 255 from Tahiti. He akua Ku,
There are four of these men; He ulele Ku mai ka lani, 595
Yes, eight of these men, He haole Ku mai Tahiti,
Ku, Lono, Kane, Kanaloa, 256 He mau kanaka ia eha.
Kanemakaiahuawahine, 600 Ewalu hoi nei kanaka,
Haihaipuaa, Kekuawalu; O Ku, O Lono, O Kane, O Kanaloa,
These are your equal. O Kanemakaiahuawahine, 600
There is Koua whose stone floor burns; O Haihaipuaa, Kekuawalu la,
The shimmering heat from the sun arises; Ua like.
Warmed are the offspring of Umulau, 605 O Kona la, ua wela ka papa,
Lanipipili, and Hooilo; Ua keehu o ka la,
The sun rises, it comes forth; Ua wela ka hua o Umulau, 605
By the power of the great-voiced Ku O Kalanipili, o Hooilo,
Was the sun given; E a’e, e puka ae ka la;
By it are warmed the selfish chiefs of Kona. 610 Ka mana o Kuleonui
Haawia mai ai ka la;
Mahana ai na ’lii aua o Kona. 610

[The following is found as a conclusion of the foregoing [Ua loaa keia malalo iho, mai a S. M. Kamakau
chant, contributed by S. M. Kamakau to the Kuokoa in ka malama o Mei, 1868, ma ke ano he panina
May, 1868, and is the “unwritten portion” referred to by ’e, a oia nohoi ka “hapa i kakau ole ia” i oleloia
C. J. Lyons in his published translation (with Judge L. o kana (ame ka Lunakanawai L. Andrews) unu
Andrews) of the Song of Kualii. 257 Fornander refers to it Kualii, i hoolahaia ’i. Ua hoakaka o Fornander
in his Polynesian Race, Vol. II, p. 284, but does not Buke Pol. Race, Buke II, aoao 284, aka, aole n
append it in his republication of the song, with a mea la me kekahi unuhina ano okoa ae, ma ka
somewhat different interpretation as an Appendix in said ana i ua Mele la, ma ke ano he Pakui no ua Bu
Volume II, from the fact that he questioned its o kona kanalua i ka oiaio o ia mea. Me ke kan
genuineness. Not doubting his judgment in the case, it manao ma keia ninau, e lilo ana ia mea maane
serves here as a further tribute, at a later period kekahi wa aku paha mahope, i kakoo ikaika i k
possibly, to the name and fame of Kualii by a historian kaulana o Kualii, ma o kekahi mea kakau moo
of his race.—Ed.] ponoi.—Luna Hooponopono.]

A CHANT FOR KUALII. HE MELE NO KUALII.

The first 258 chief, the second chief, Akahi kalani, alua kalani,
The third chief, the fourth chief, Akolu kalani, aha kalani,
The fifth chief, the sixth chief, Alima kalani, aono kalani,
The seventh chief, the eighth chief, Ahiku kalani, awalu kalani,
The ninth chief, the tenth chief is Ku, 5 Aiwa kalani, umi kalani ia Ku. 5
Kuikealaikauaokalani 259 encircled. Ia Kuikealaikauaokalani, a puni.
The first warrior, the second warrior, Akahi koa, alua koa,
The third warrior, the fourth warrior, Akolu koa, aha koa,
The fifth warrior, the sixth warrior, Alima koa, aono koa,
The seventh warrior, the eighth warrior, 10 Ahiku koa, awalu koa, 10
The ninth warrior, the tenth warrior Aiwa koa, umi koa,
Is Kalanikahimakaialii, 260 Ia Kalanikahimakaialii,
The young warrior of all Maui. I ke keiki koa o Maui a puni,
Rent is Kauakahi, 261 one; Owa Kauakahi-akahi,
The second is mistaken, 262 two; 15 Kuihewa i ka lua-alua, 15
Kaihikapu is third, three; He kolu Kaihikapu-akolu,
Kukawelo is fourth, four; He ha Kukawelo-aha,
Lononuiakea 263 is fifth, five; Lononuiakea i ka lima-alima,
Kahoowaha the sixth, six; He ono Kahoowaha-aono,
The seventh is at Kapua, seven; 20 He hiku ai Kapua-ahiku, 20
Hoalani is [an] even [number], even; He like Hoalani-a like,
The sea beach of Hua is ninth, He iwa kahakai o Hua,
The sea beach where I was wont to ramble. Kuukahakai e paiai a puni
Kalani has encircled Kahiki, Puni Kahiki ia Kalani.
The flying of the great koae, 264 25 I ka lele a ke koae nui, 25
The koae that soars high above A ke koae lele kau maluna.
Is Kapiiohookalani. 265 O Kapiiohookala—ni,
You have ruled 266 the land [396] Ua ai oe i ka aina [397]
Of Waialua, of Waimea, O Waialua, O Waimea,
Places endeared to the chief. 30 O na wai aloha a Kalani, 30
Kupihea 267 has fled Ua hee Kupihea,
And is out on the ocean, Aia i kai i ka moana,
The eye looking on Oahu. Nana ka maka i Oahu.
Like the entrance to a house Me he ipuka no ka hale la,
Is the ravine 268 of Pohakea, 35 Ke kowa o Pohakea 35
Maunalahilahi standing beside. Ke ku a Maunalahilahi,
The sailing to Kanilio, Ke holo ana i Kaneilio.
Moving on, yes, moving on, Panee la-e-panee,
Moving on to Waihee. Panee aku a Waihee,
Landing at Kapahukukui 40 Pae no i Kapahukukui, 40
[He] seized and hurled the weapons around. Hopu no pahu i ka laau a puni;
Turning, yes, turning, Hookaa-la-e-hookaa
Turning the canoes seaward Hookaa na waa makai,
You land at Maeaea. Kau oe i Maeaea.
Kaweloiki of Aila gave 45 Haawi o Kaweloiki a Aila, 45
The food of Kamananui, O na ai o Kamananui,
The food of Palaa. O na ai o Palaa.
Your fame resounds in Nahuina, Kani ko pihe i Nahuina,
’Tis the fault of Kanealala, Kihewa a Kanealala.
The son of Kaio the elder, 50 O keiki a Kaiao nui, 50
The pig for the king, O ka puaa i ka Moi,
The lord that craves for the goose. O ka haku-ne i ka nene,
The uau 269 screams, agitated, disturbed; U o ka ua’u, li-o o ka li-o,
The ao calls as the rod is poked in; 270 A o ka ao, hou ka houhou,
When pressed, yea, it shrinks. 55 Ke anunu, ka ekekekeiau-e, 55
When the mice 271 glance aside the owl is gone, Awihi ka iole, heo ka pueo,
The auku 272 cries, the duck extends its neck. Owa ka aukuu, aeae Koloa,
At Waikoloa in Keawewaihe O Waikoloa ia Keawewaihe,
[At the] mudstone the birds cluster Pohakulepo pae mai ka manu,
Within the depression. 60 Iloko o ka poopoo, 60
The ao, the ao, the agitated, the agitated, Ka ao ka ao-e—, ka li-o— ka lio-e
Going over the land in flocks, Kakakai honua, puaoao,
They ate at the boundary of Kaakiko stream. Ai nui, pale kahawai kaakiko e,
The first drawing of Ewa’s net Ka makauakahi o Ewa,
Entrapped the fish of Mokumoa; 65 Ua puni ka ia o Mokumoa, 65
They are strewn on the grass. Ua kau ia i ka nene,
The kalo danced, danced noisily, Ua haa kalo, haa nu,
The fish of Kawelo danced, Haa ka ia o Kawelo,
The potatoes of Pahua danced, Haa na uala o Pahua,
The mahiki grass at Puukea danced, 70 Haa ka mahiki i Puukea, 70
The ununu 273 danced at Peleula, Haa ka ununu i Peleula,
Makaaho danced on the way. 274 Haa Makaaho i ke ala,
Say, Ku, the rain comes by way of Kekaha, Ku, E Ku-e-ma Kekaha ka ua e Ku,
Bringing food 275 for the fish of Maunalua; I ai na ka ia o Maunalua.
The rain that supplies Kekuapololi, 75 Ua nana ia Kekuapololi, 75
Coming near to the hill of Kalalau Hoae ia i Puuokalalau,
Koolau trembles. Naueue Koolau,
O Ku, O Kalamahaaiakea, 276 E Ku e Kalamahaaiakea,
To Kalama indeed that land belongs, No Kalama no ia aiua,
A permanent resident in Kailua. 80 He aina noho mau i Kailua, 80
Waimanalo commences to be jocular Hoomaka lealea aku Waimanalo,
While they oppose Kaneohe. 277 E kii e peku o Kaneohe.
Yea, from Mahinui to Haakalo Mai Mahinui a Haakalo-e
Until Kuikealaikauaokalani 278 reigns, A ai o Kuikealaikauaokalani,
At the back, in front, and all around. 279 85 Ma ke kua, ma ke alo a puni 85
To Kahiki of Paumakua the chief, A Kahiki ia Paumakuakalani,
To Kahiki in Keakeaau the chief. A Kahiki i Keakeaau o ka lani,
By Wakea the chief, Kahiki is sheltered. Ia Wakea ka lani, a malu Kahiki,
Peaceful is Kahiki. Malumalu Kahiki.
In the rainy season of Maheleana, yea, 90 I ke ao ua o Maheleana-e— 90
Mullet is plentiful at Kauai. Amaama ka Kauai
It reaches to Puna and outward Ua hiki ma Puna mawaho,
Even to Kau, to Hilo. Ma Kau, i Hilo,
The land is seen to be peaceful Ua ikea ka like o ka aina,
By a certain chief, offspring of the heaven, 95 E kekahi alii hua o ka lani, 95
By Keakaliilani of Waipa. A Keakaliilani o Waipa,
Lest you will be overtaken, Kipu, mischief-maker of O loohia oe e Kipu, ahiahi o Palaa
Palaau, E ke kiu ahiu ilalo o Kahana,
By the wild spy 280 below at Kahana Hoolawa i kekahi alii,
For supporting a certain chief. E puni e Kalani-e-ko aina-e— 100
Say, chief, reign over your land, 100 A Kaiwi i Kekuapolo-li—
From Kaiwi to Kekuapololi A ka wai i ke hoomanalo,
To the water that quenches not, Ke kai kalua i ka manawa,
The sea that marks the seasons, Ka ohe oki i ka ili, [399]
The bambu that cuts the skin [398] O Ka maka o ka ua o Haao, 105
Of the eye 281 of the rain of Haao, 282 105 A ka hee i ka ua alaea o Kii,
The retreating red 283 rain of Kii, A ke kanaka luu hala o Halakua,
Of the pandanus diver of Halakua, 284 Ka wai i ka hee moana
Of the waters of the ebbing tide, Ka ai ka hoea ana
And the food that is served on arrival. Ka wai hole a ka ili 110
The water that cleanses the skin 110 I ka wai e hookane ana,
[Is] the water which befriends a man Ka hakipuu i ka manawa,
At the change of the season, Ka loana o ka aina
The length and breadth of the land. Ka awa loha i ka ili
The awa that withers the skin O ka maka i ka ua o Haao. 115
Of the eye by the rain of Haao. 115 A ka hana i ke ka wailehua o Hilo,
As one labors drawing the nectar of the lehua 285 A ka luuluu i ke hoaloha,
blossoms of Hilo, A ka puhee miki i kai
Like burden to a friend, A ka aha i ke hili a paa
Is the arm of the squid that draws seaward, O ka paa i ka hoolua, 120
Like cords braided tightly O ka hoolua i ka hoonui
Like the grip of the summer wind 120 Hiki ka pona i o honau,
That increases in strength O ma i ka hookaona,
Until its fury reaches the end O ke kaa ma i ka wela
And ceases in its anger. O kai a Alapai inu a maloo 125
Of the heat that rolls on; Papau kai e Koholalele,
Of the sea that Alapai drank up dry. 286 125 O Malae i ka ie waa,
Shallow is the sea of Koholalele; Me ka ie waa i kai,
Of Malae where the canoe-vines 287 grow, O Malae i ka hookahana,
Where the canoe-vines grow near the sea I ka pae puu e loaa, 130
Of Malae, the little cove O Keana kai akea,
Bounded by a cluster of hills. 130 O ka hoohukuhuku i ka hala lei,
At Keana [of the] wide sea, I ka hala lei o Puna.
Where the hala wreaths are piled up, Uaua hiki kona, iluna ka malie,
The hala wreaths of Puna. Ke hele ae la e Kuhoopepelaualan
The south winds have arrived, the upper lands are dry; Hele ae o Kaipunui ko aina-e—
There travels Kuhoopepelaualani, 288 135 A ka opana i opanaiki
Traveling on Kaipuni, your land. A ka oio e hana, e lomia mai la,
Holo ka oo alahee a ka mahiai,
At the cutting 289 of the pieces
The oio 290 is prepared, being mashed there. E mahiai mai la, 140
Speedy is the alahee 291 digger of the cultivator Holo ka o i ke kia manu,
As he toils there. 140 Ka pili apane o ka nahele
The bird-catcher is speedy with his trap-pole, Ua wele i ka la na mauu pili
Catching the apane 292 of the woods. I ka la o Makalii,
The pili-grass is scorched by the sun, A ka pahukauila i ke kanaka pahip
The sun of Makalii 293 [season]. O Wailoa ia i na hoa i ka olelo
The kauila drum 294 of the fun maker, 295 145 O ka hookauna i ka la o lalo he m
Is Wailoa, so say his companions. A ka pupu o ka awa ua o Mahelea
As the sun declines it is calm. A ka wai i ke hoea ’na
With the drinking of the awa the rain fell at Maheleana, A ke kukui o Hapuu ke hele ia, 15
The waters appeared, O ka pupu e loa ke pae ia
And the kukui trees of Hapuu, they moved. 150 O ka punaunu inai o ka nenue
Like a strange 296 shell that drifts ashore O kuu palauhu wauke
Is a low-born who partakes with a chief. Alai hoihoi o ka makani
My bundle of pulpy wauke, 297 A ka wai i ke hooloa 155
Obstructed, yet returned by the wind, A ka pa i na muliwai
So is the water in the trench, 155 I ka mana kai e nui,
Even to the wall by the river, Hina kokea i na lua wai
At the main branch that flows to the sea. Malino o Hikiau
White cane fell at the wells, A Kailikahi i ka ili hoaaloha, 160
Calm is the sea of Hikiau Ka aukuu i o Mokupaoa,
At Kailikahi, the friendly district, 298 160 Kawaihaipai lehua o ka holo.
The aukuu of Mokupaoa. A Kealia manu o Anahola [401]
The lehua of Kawaihaipai runs A keena i ke ahu a Kupihea
To the bird swamps of Anahola, [400] I kona au. 165
A nook near the heap of stones 299 of Kupihea Hele ae o Kaipunui ko aina-e—
In his time. 165 A Kaula i ka pali ua o Hanalei.
A ka ha o ka leo
Move on to Kaipuni, your land; I ka aukai o lalo he malie,
To Kaula, the rain-swept cliffs of Hanalei. A ka ohiki haehae kapa o Waiolon
When the voice is hoarse A ke kanaka kaha nalu o Makawa
Of the swimmer below in the calm, Hihi na maile, hihi i kai o Kua,
[Where lives] the cloth-tearing sand-crab 300 of Waiolono, A ka wai kulu i ka wai hoanae
170 A ka maili i ka ohuku,
And the surf rider of Makawa. O ka ohuku i ke hoolae. 175
The maile 301 vines creep, creep down to Kua, A ka hawahawa i ka pili a ka ua
To the dripping water, the sacred water O ka ua pili o Hooilo-e-a,
At the pebbly beach at the knoll; O Kuikealaikauaokalani no ke ’lii.
The knoll at the small point, 175
Where, drenched by the heavy rains, O Kawelo-e, e Kawelo-e,
Yea, the heavy rains of winter, 302 O Kaweloiki puu oioi, 180
Kuikealaikauaokalani is indeed the king. Puu o Kapolei-e—
Uliuli ka poi e piha nei—o Honouli
O Kawelo! Say, Kawelo! Aeae ka paakai o Kahuaiki—Hoae
Kaweloiki, the sharp-pointed hill, 180 Pikele ka ia e Waikele—o Waikele
Hill of Kapolei. Ka hale pio i Kauamoa—o Waipio;
Blue is the poi which appeases [the hunger] of E kuu kaua i ka loko awa—o Waia
Honouliuli; Mai hoomanana ia oe—o Manana
Fine the salt of Kahuaike—Hoaeae; He kini kahawai,
Slippery is the fish of Waikele—Waikele; He lau kamano—o Waimano;
The arched house at Kauamoa—Waipio; 185 Ko ia kaua e ke au—o Waiau; 190
Let us cast the net in the awa-pond—of Waiawa; Kukui malumalu kaua—Waimalu;
Do not stretch yourself at—Manana. E ala kaua ua ao-e—o Kalauao;
Many 303 are the ravines, E kipi kaua e ai—o Aiea;
Numerous 304 the sharks, at Waimano; Mai hoohalawa ia oe—o Halawa;
We are drawn by the current of Waiau; 190 E noho kaua i ka lua—o Moanalua
In the kukui grove we are sheltered—in Waimalu; Hoopiopio hau kaua—o Kahauiki;
Let us arise, it is daylight—at Kalauao; Hookeekee lihi kaua—o Kalihi;
Let us enter and dine—at Aiea; E pii kaua i ka lama—o Kapalama
Do not pass by—Halawa; E nunu a paa hoawe—o Hononun
Let us abide in the hollow—of Moanalua; 195 Kiki kuu oho ilaila—o Waikiki; 200
We will bend the hau—at Kahauiki; Kike ka hua o ka alae—o Waialae;
And go zigzagging down the edge—of Kalihi; He wahine oho lupe keia—o Wailu
Let us go up for lama 305—in Kapalama; E pii kaua i niu—o Niu;
Then bundle and fasten on the back—at Hononunu; He wahine heekoko keia—o Koko;
There my hair is anointed—at Waikiki; 200 Ouou ka manu o Kaula—o Kuliouo
The egg of the mud-hen 306 is broken—at Waialae; Ua puni ka moku ia Ku,
This is a woman with flowing hair—Wailupe; Me he kai la, me he wai la,
Let us climb to get coconuts—at Niu; Me he kai malu la ka hoi;
This is a woman with catamenia—Koko; Ka poe kou i kai e Kahaloa,
The bird of Kaula is singing—at Kuliouou; 307 205 Ku ke koa, ku ka o’a, 210
The island is encompassed by Ku, Ku ka lehua, ku ke alii,
As by the sea, as by the water, Ku ka hele a maua.
As a sheltered sea indeed; O Kuikealaikauaokalani,
The kou trees down at Kahaloa, A puni—Amama—ua noa.
The koa stands, the o’a stands, 210
The lehua stands, the chief stands.
We can wend our way
With Kuikealaikauaokalani,
Compassed. It is opened. 308 It is free.

At the conclusion of the chanting of the mele by A pau keia pule ana a Kapaahulani i keia pule,
Kapaahulani, the two armies came together and the kaua elua, a hoomoe wale ke kaua. Ia manaw
battle was declared off. The king of Koolauloa then gave alii o Koolauloa i kona mau okana Koolauloa, K
over, or ceded, the districts of Koolauloa, Koolaupoko, Waialua, a me Waianae. A lohe ko Kauai alii i k
Waialua and Waianae. When the king of Kauai heard Kualii akamai i ke kaua, a me kona lanakila an
how Kualii excelled over all the others in war, and how kaua ana ma Honouliuli i kula o Keahumoa, ia
he had gained the victory at the battle of Honouliuli on la ua alii nei o Kauai e halawai me Kualii, a haa
the plains of Keahumoa, he came to meet Kualii and Kauai me Kualii, a lilo ae la na moku a pau ia K
gave him Kauai, and by this act Kualii became possessed Niihau. [403]
of all the islands from Hawaii to Niihau. [402]

CHAPTER III. MOKUNA III.

 KAPAAHULANI AND HIS BROTHER KAPAAHULANI A ME KONA KA
KAMAKAAULANI. KAMAKAAULANI.

After the battle of the plains of Keahumoa as related in Mahope iho o ka hoouka kaua ana i ke kula o
the preceding chapters, Kapaahulani, the elder brother me ka mea i oleloia ma na Mokuna mua, a o K
of Kamakaaulani, the one on the side of the king of kaikuaana o Kamakaaulani ka mea ma kela ao
Koolau, received great riches from the king. Kapaahulani mai. Hooili ia aku la ka pomaikai a me ka waiw
also became a great favorite with the king, and the king lilo ae la oia i punahele nui na ke alii; a haawi
gave away his own lands to him; and on the same day mau aina ponoi no ua Kapaahulani nei, a ma k
in which the battle ended and while on their return by kaua ana, ia lakou i hoi ai a uka o Lihue, hook
way of Lihue, Kapaahulani was presented with swine, ka ai, a me ka waiwai; a haule lakou nei i lalo
food and various other things; and when they arrived at hookupu no ka waiwai.
Waianae presents of property were again given him.

It was while on this return to Koolau that Kapaahulani Ia manawa, olelo aku la o Kapaahulani i ke ali
remarked to the chief these words: “Here I am waiwai, ke noho la ko’u kaikaina a me ka’u wa
possessed of all this property, while my younger brother mai i ko’u puu waiwai.” A no keia olelo a Kapa
and my wife are without any knowledge of what amount ae la ka pukaua o ke alii i mau waa, mamuli o
of property I am in possession of.” e lawe i ka waiwai hookupu na ko Kapaahulan
laweia ka waiwai maluna o na waa kaulua, a lo
Because of these words spoken by Kapaahulani, the ma Puuloa, a malaila i haawiia aku ai ka waiwa
chief warrior of the king, in obedience to the order of
the king, picked out a certain number of canoes and
loaded them with some of the presents given to
Kapaahulani to be taken to his people. After the
presents were loaded into several large double canoes,
they were sent out to meet Kamakaaulani who was then
at Puuloa, and there the presents were given him.

Before Kamakaaulani parted from his brother they had A o Kamakaaulani hoi, mamua o ko laua hook
an understanding where he would be at the close of ua kuka mua no laua i kahi e noho ai o Kamak
hostilities. This was why Kamakaaulani resided at iho o ka hoomoe ana o ke kaua; a nolaila no i
Puuloa, that being the place agreed upon by them. Thus Kamakaaulani ma Puuloa, mamuli o ka laua ku
by this conduct of theirs both sides were deceived keia hana a laua ua puni na aoao elua ia laua
through their duplicity. nolu ana.

about the return of the king to waialua. no ka hoi ana o ke alii i wai

When Kapaahulani was returning to Waialua with the Ia Kapaahulani i hoi ai me ke alii i Waialua ma
king, after leaving Waianae, the king said to him: “How haalele ana ia Waianae, i aku la ke alii ia Kapa
about the lands I gave you while we were on the plains aina a’u i haawi aku ai ia oe i ka la i hoomoe a
of Keahumoa on the day the battle was called off?” Keahumoa?” I aku la o Kapaahulani: “E ke alii,
aina, aole no hoi kekahi mea e ae. Aka, o ka’u
Kapaahulani replied: “Listen, O King! I do not care for ai he pono i ke alii ke haawi mai, o ka halepap
the lands nor for anything else. The only thing I want alii ua pono ka Kapaahulani olelo ma ia mea, a
you to give me is to have the general care of your store Kapaahulani i puuku nui no ka halepapaa, ma
houses.” 309 When the king saw that Kapaahulani had
made the proper answer in the matter, he was
thereupon made chief steward over the store houses in
the name of the king.

After he became chief steward and had control of the Alaila, o ka hana a ua Kapaahulani nei, iaia ma
king’s store houses, he took it upon himself as a duty to no ka halepapaa, o ka hoouna mau i na mea a
continuously supply his younger brother with the halepapaa no kona kaikaina no Kamakaaulani.
different good things in the store houses, in accordance ana o ka laua mea i manao mua ai. [405]
with their first agreement. [404]

CHAPTER IV. MOKUNA IV.

THE GENEALOGICAL TREE OF KUALII KA PAPA KUAUHAU O KUALII, MAI

FROM KANE TO WAKEA. HIKI IA WAKEA.—Contin

HUSBAND. KANE. WIFE. WAHINE. CHILD. KEIKI. HUSBAND. KANE. WIFE. WAHINE.
Kahai. Hinauluohia. W
Kane. Hulihonua.
Wahieloa. Koolaukahili.
Kanaloa. Laka. Laka. Hikawaolena.
Ukinaopiopio. Luanuu. Kapokulaiula.
Kauakahi. Kamooalewa.
Kamea. Popomaili.
Maliu. Maluakapo. Pohukaina. Huahuakapolei.
Hua. Hikimolulolea.
Hulihonua. Keakahulilani. Laka.
Pau. Kapohaakia.
Laka. Kapapaiakele. Maluakapo.
Kinilauaemano. Ulupalu. Halo. Kapoea.
Huanuikalalailai.
Halo. Koniewalu. Kamanonookalani.
Molehai.
Kamanonookalani. Kaehuaokalani. Keohokalani.
Keohokalani. Kaamookalani. Kaleiokalani. Paumakua. Manokalililani.
Kaleiokalani. Kaopuahihi. Kalalii. Haho. Kauilaianapa.
Kalalii. Keaomele. Haule.
Haule. Loaa. Nanea. Polena. Hikawainui.
Nanea. Walea. Nananuu.
Nananuu. Laholana. Lalokona. Hanalaanui. Mahuia.
Lalokona. Lalohoaniani. Hanuapoiluna.
Lanakawai. Kalohialiiokawai.
Hanuapoiluna. Hanuapoilalo. Pokinikini.
Laau. Kukamolimolialoha.
Pokinikini. Polehulehu. Pomanomano. Pili. Hinaaiaku.
Pomanomano. Pohakoikoi. Kupukupunuu.
Koa. Hinaaumai.
Kupukupunuu. Kupukupulani. Kamoleokahonua.
Ole. Hinamailelii.
Kamoleokahonua. Keaaokahonua. Ohemoku. Kukohou. Hinakeuki.
Ohemoku. Pinainai. Mahulu.
Kaniuhi. Hiliamakani.
Mahulu. Hiona. Milipomea.
Milipomea. Hanahanaiau. Hookumukapo. Hualani.
Kanipahu.
Hookumukapo. Hoaono. Luakahakona.
Alaikauakoko.
Luakahakona. Ninau. Kahiko.
Kahiko. Kapulanakehau. Wakea. Kalapana. Makeamalamaihanae.
 HUSBAND. KANE. WIFE. WAHINE. CHILD. KEIKI. HUSBAND. KANE. WIFE. WAHINE.
Wakea. Papa. Hoohokukalani. Kahiamoeleaikaaikupou. Kapoakaulahailaa.
Wakea. Hoohokukalani. Haloa. Kalaunuiohua. Kaheka.
Haloa. Hinamanouluae. Waia.
Waia. Huhune. Hinanalo.
Hinanalo. Haunuu. Nanakehili. Kuaiwa. Kamuleilani.
Nanakehili. Haulani. Wailoa.
Wailoa. Hikawaopuaianea. Kio.
Kio. Kamole. Ole. Kahoukapu. Laakapu.
Ole. Kai. Pupue. Kauholanuimahu. Neula.
Pupue. Kamahele. Manaku. Kiha. Waoilea.
Manaku. Hikohaale. Kahiko.
Liloa. Pinea.
Kahiko. Kaea. Luanuu.
Luanuu. Kawaamaukele. Kii. Akahiakuleana.
Ulu. Kulamea.
Kii. Hinakoula. Makaalua.
Nanaulu.

Nanaulu. Ulukou. Nanamea.

Nana. Kapukini.
Umi.
Ulu. Kapunuu. Kapulani.

Nanaiea. A
Nanaiea. Kahaumokuleia. Nanailani. Piikea.
Nanailani. Hinakinau. Waikulani.
Waikulani. Kekauilani. Kuheleimoana. Keliiokaloa. Makuwahineopalaka.
Kuheleimoana. Mapunaiaala. Konohiki.
Konohiki. Hikaululena. Wawena.
Kukailani. Kaohukiokalani.
Wawena. Hinamahuia. Akalana.

Mauimua.
Makakaualii. Kapukamola.
Mauihope. Keawenuiaumi. Koihalawai.
Akalana. Hinakawea.
Mauikiikii.
Kanaloakuaana. Kaikilani.
Mauiakalana.

Mauiakalana. Hinakealohaila. Nanamaoa.

Nanamaoa. Hinaikapaekua. Nanakulei. Keakealanikane. Kealiiokalani.
Nanakulei. Kahaukuhonua. Nanakaoko. Iwikauikaua. Keakamahana.
Nanakaoko. Kahihiokalani. Heleipawa. Kanaloaakapulehu. Keakealani.
Heleipawa. Kookookumaikalani. Hulumalailani. Kaneikauaiwilani. Keakealani.
Hulumalailani. Hinamaikalani. Aikane.

Puna. Keawe. Kalanikauleleaiwi.

Aikane. Hinahanaiakamalama.
Hema.
Keeaumoku. Kamakaimoku.
Puna. Hainalau. Ua. Kekela. Haae.
Hema. Ulamahahoa. Kahai. Kalanikupuapaikalaninui. Kekuaipoiwa.
[406] [407]
The genealogical tree from Kane to Kahiko contains O ka papa mookuauhau mai a Kane mai a hiki
twenty-nine generations and from Wakea to iwakaluakumamaiwa hanauna; a mai a Wakea
Kamehameha seventy-five generations. By adding the Kamehameha, he kanahikukumamalima hanau
generations before Wakea with those after him we have ka hanauna mamua aku o Wakea, hookahi ha
one hundred and fourteen generations. We cannot, hanauna. Aka, aole nae e hiki ke hoomaopopo
however, ascertain whether this is so, nor can we say pololei keia mau papa mookuauhau. Pela nae
that this genealogical tree is correct. This, however, is moolelo o Kualii.
what is shown in the history of Kualii.

There are several genealogical trees in connection with He nui na mookuauhau e pili ana i keia mooku
this genealogy of kings; the following is one of them: ka papa kuauhau malalo:

HUSBAND. WIFE. CHILD. KANE. WAHINE.

Kapapaiakea. Kauhihi. Hinakapeau. O Kapapaiakea. O Kauhihi. Hin
Hinakapeau. Ukunohunohu. Ukunaopiopi. O Hinakapeau. O Kunohunohu. Uk
Ukunaopiopi. Maakuanana. Kalei. O Ukunaopiopi. O Maakuanana. Kal

Kaiakea. Kai

Kalei. Kaeelekoha. Moanakea. Kalei. O Kaeelekoha. Mo

Hulukeeaea. Hu

Hauii. Ha
Kaiakea. Kaehokumanawa. Kaiakea. Kaehokumanawa.
Hauee. Ha

Moanakea. Kauakahikuaana. Kanehoalani. Moanakea. Kauakahikuaana. Kan

Iwikauikauanui. Kauakahikuaana. Hauonunaholoholo. Iwikauikauanui. Kauakahikuaana. Ha
Hulekeeaea. Kahakuakea. Hauiikaiapokahi. Hulukeeaea. Kahakuakea. Ha

Uliuli. Uliu
Hauiikaiapokahi. Wahineikapeakapu. Hauiikaiapokahi. Wahineikapeakapu.
Maihea. Ma

Uliuli. Kaukeano. Uliuli. Kau

Kahakapolani. Kahakapolani.
Maihea. Mehameha. Maihea. Me

Uliuli. Niau. Kahiko. Uliuli. Niau. Kah

Kahiko. Kupulanakehau. Wakea. Kahiko. Kupulanakehau. Wa

In this genealogical tree there are eleven generations Ma keia papa kuauhau, he umikumamakahi ha
from Kapapaiakea 310 to Wakea. It is, however, told that Kapapaiakea mai a hiki ia Wakea. Ua oleloia n
the genealogical tree to which Kane is the head, and the mookuauhau o Kane, a me ka papa kuauhau o
genealogical tree of Kapapaiakea, were handed down by ko Oahu nei poe malama kuauhau ia, a ua ike
those who had the keeping of the Oahu genealogy, and elua ma ka moolelo o Kualii. A o ka mookuauh
these divisions are seen in the history of Kualii; and the a hiki ia Wakea, mai a Wakea a hiki ia Kameha
genealogical tree from Opuukahonua 311 to Wakea and poe malama kuauhau ia. A ua ike ia pela ia pa
from Wakea to Kamehameha had been handed down by ka moolelo o Moikeha. Aka o ka mea i ike mau
the Hawaii genealogy keepers, and this genealogical o ka mookuauhau mai a Wakea mai a hiki ia K
tree is seen in the history of Moikeha. But the mamuli o ka mea i olelo ia e Kalauwalu, a me
genealogical tree that is commonly seen these days is kuauhau e ae, a ma ka hoomaopopo ana i ka
mookuauhau, aole he akaka; aka, ma ka nana
the one from Wakea to Kamehameha as told by ana o na mookuauhau me he mea la, ua kaaw
Kalauwalu and other genealogy keepers. mookuauhau, a kaawale ko Kauai. Aka, o ko M
he okoa. Ma ka malama ana a ko Molokai poe
In trying to ascertain the truth of the different divisions o Hookumukahonua ke kupuna mua o ko Haw
of these genealogical trees one is left in doubt as to Aka, ma ka manao ana o ka poe kakaolelo, ma
their correctness, but in looking them over one cannot ka hookumu ana o ka hanauna alii.
help seeing that each island had a separate tree, the
Maui one being different from that of Kauai. Molokai’s
genealogy differs again. In the records kept by the
Molokai genealogist it is stated that Hookumukahonua
was the progenitor of the royal family of Hawaii, but in
the opinion of historians they generated from Wakea.

CHAPTER V. MOKUNA V.

THE BATTLES OF KUALII AND THE BATTLE NA HOOUKA KAUA A KUALII, A M

GROUNDS. KAUA.

The battle on the plains of Keahumoa at Honouliuli, Ua oleloia ma ka Mokuna I ka hoouka kaua an
Ewa, is described in Chapter I. In looking over the Keahumoa, i Honouliuli, ma Ewa, a i ka nana a
history of Kualii related in that chapter, it is thought to Kualii ma kela mokuna, ua manaoia oia ka hoo
have been the last battle in which he took part, for in a Kualii. No ka mea, ma ka Mokuna II, ua olel
Chapter II it is shown that Kauai was simply given over ia mai o Kauai e ko Kauai alii ia Kualii; nolaila,
by its king to Kualii, whereby the whole group from aina mai Hawaii a Niihau. Pela i manao ia ai, o
Hawaii to Niihau was united [under him]. Therefore it is a Kualii. [409]
believed that was Kualii’s final contest. [408]

The first battle on Oahu in which Kualii took part where O ka hoouka kaua mua a Kualii i hoomaka ai k
a general war was had, was the one fought on Oahu, oia ka hoouka kaua ana iluna o Kawalu
Kawaluna, the heights above Waolani, 312 where a great Waolani, oia ka luku nui ana i ula pu ai ke pili
slaughter took place that reddened the pili grass of penei ka moolelo o ia kaua ana:
Keanakamano. The history of that battle is told as
follows:

Oahu had four kings just prior to the time of Eha mau alii o Oahu mamua aku o Kakuhihew
Kakuhihewa; Lonohulimoku was the king of Koolaupoko; oia ke alii o Koolaupoko; a o Lonohulilani, oia
Lonohulilani was the king of Koolauloa and Waialua; me Waialua; o Lonokukaelekoa, oia ke alii o W
Lonokukaelekoa was the king of Waianae and Ewa; and Lonoikaika, oia ke alii o Kona, mai Moanalua a
Lonoikaika was the king of Kona, from Moanalua to
Maunalua. While Kualii was residing at Kalehuawehe, in Aka, o Kualii i kona manawa ma Waikiki, iaia e
Waikiki, at a time when he was about to attain the age Kalehuawehe, iloko o kona mau la e hookanak
of manhood, he began to be dissatisfied with the king of pono ole iho la kona manao i ko Kona nei alii;
Kona district, because his immediate attendants often kahu ponoi o Kualii, ua kaniuhu mau lakou no
A no ia mea, hele aku la ua mau kahu nei o Ku
olelo aku la, me ka i aku: “Ina no hoi paha ka
complained of being oppressed and would come to him a he waawaa ikaika i ke kaua, ola nei mau iwi
with the following remarks: huhewa; kai ka lilo o ka hoounauna ino i na ’li
ko wa kamalii ka hoi, paeha oe, kai no ko koa
“If your muscular body was only that of a fearless mau la ia koa ou. Eia ka he koa ia no ka wa ka
warrior these bones would indeed be saved: but no, Kualii: “Ua kaua, ua loaa mai la ia oukou ka hu
your strength is worthless. Here we are being ordered mau la koe ula ke pili.”
roughly by the different chiefs which is so degrading and
angers us. In your younger days you could beat
everybody whom you fought against. Being so fearless
in your childhood days, one would think it would
continue; yet alas, it was only the fearlessness of youth.”
Kualii replied: “There will be fighting then, since you
have found the cause why I should urge it. A few days
hence the pili grass will be reddened.” 313

On the expiration of the days during which the temple A pau na la o ke kapu heiau iluna o Kawaluna,
on Kawaluna was dedicated, 314 the following night the aku la ke kaua a Lonoikaika iluna o Keanakam
army of Lonoikaika arrived on Keanakamano, as word oleloia e Lonoikaika, ua kipi o Kualii. A oia no
had been carried to Lonoikaika that “Kualii has rebelled.” ai na koa ma ke kula o Keanakamano. No ka m
This was the reason why the soldiers slept that night on ana ia Kawaluna, aole i kupono ia ia Kualii.
the plains of Keanakamano, Kualii in dedicating the
temple on Kawaluna had overstepped himself. Very early Ma ka pili o ke ao, hoala aku la o Kualii i kona
that morning Kualii aroused his father Kauakahiakahoowaha: “E! Auhea oe, hoalaia a
Kauakahiakahoowaha 315 with the words: “Say, Where art ua puni kakou i ke kaua, eia malalo mai o kako
thou? Rouse up the men, we are now surrounded by the eia ma Koolau mai kekahi kaua, a eia malalo o
enemy; there is one army below us, there is another kaua; hookahi wale no pali i koe o Waolani; no
army from Koolau and there is still another one from oe i na kanaka, a e hoakoakoa mai, no ka mea
Waialua; there is but one pali left, that of Waolani, no ke kaua.”
therefore you must rouse up the men and get them
I aku la o Kauakahiakahoowaha: “Pehea i mao
together as I am ready for the battle.”
puni kakou i ke kaua?” I aku o Kualii: “Ke hai m
Kauakahiakahoowaha replied: “How do you know that
kaua ko ke ao, no ka mea, ua olelo ae la ke al
we are surrounded by the enemy?”
kipi kakou, no ko kakou kii ana mai nei e kapu
Kualii spoke up: “The night tells me that there will be nei. No ka mea, he akua nui ka mea nana e ka
war in the day time, for the king, Lonoikaika, has
I aku o Kauakahiakahoowaha: “Auhea oe e Ku
remarked, that we have rebelled against him, because
hai mai la ka po ia oe he kaua ko ke ao ana ae
we have come here to dedicate this temple on
mai nei hoi oe, hookahi pali i koe aole i paa i k
Kawaluna, thus taking upon ourselves something which
a eia ko’u manao, ma ia pali kakou e holo aku
only a great god has power to do.”
hiki aku ma Waikiki.” I aku la o Kualii: “Heaha
Kauakahiakahoowaha replied: “Say, Kualii, since the
ai, a holo aku auanei pakele ina he kaua no ke
night has told you that there will be war during the day
make; e holo no paha a he make, e make ana
and you say there is left us but one more pali, that of
oia ko kakau, e ola ana no.” [411]
Waolani, my idea is this: let us escape by way of that
pali this early morning and return to Waikiki.” Kualii
replied: “Why should we run? Do you suppose that we
would be saved by escaping? If we are to die in this
battle, running will not save us, we would indeed die;
and if we are to live, we will surely live.” [410]
Kauakahiakahoowaha again asked: “What are we to do I aku la o Kauakahiakahoowaha, me ka ninau
then?” Kualii replied: “Let us remain and fight them.” kakou?” I aku la o Kualii: “E noho kakou e kau
Kauakahiakahoowaha remarked: “If you want to fight, Kauakahaikahoowaha: “Ina i makemake oe i k
you may do so, but as for myself I am going to look for oe; aka, owau nei la, e imi ana wau i ko’u wah
a way of escaping.” Kualii then said: “You must not go; o Kualii: “Mai hele oe, noho iho pela, ina e hel
remain where you are; if you go, I may not be able to auanei wau e ike aku ia oe, make e mai oe i k
see you, for you might get killed by mistake; it is best aho no e noho mai oe a make pu iloko o ke ka
that you stay with me and let us die together in this
battle against Lonoikaika if need be.”

This conversation with his father took up a good part of A no ia mea, ua lilo loa ko Kualii manawa i ke
Kualii’s time and the day grew brighter. When it became makuakane ma keia mea a malamalama loa. I
broad daylight, Kualii looked forth and behold the pili aku auanei ka hana o Kualii, he ula pu wale la
grass was red with men; the pili grass of Keanakamano pili, ua pani ia iho ke pili o Keanakamano paap
was entirely covered with men. Kualii at this time manawa, hiamoe iho la o Kualii, aole nae he h
covered himself over as though asleep; he was not, mea e maopopo ai i kona makuakane a me ko
however, really asleep, but he did this to show his father ua pololei ka olelo a Kualii ia wanaao. I na kan
and their men that he had indeed spoken the truth that Kauakahiakahoowaha e hiamoe ana, lohe ae la
early morning. While the men and Kauakahiakahoowaha uwauwa ma kuahiwi mai, ma kahi e kokoke m
were sleeping they heard a great commotion from the Nolaila, hikilele ae la o Kauakahiakahoowaha,
mountain, somewhere near Kawaluna. ka hana, e hoonoho ana ke kaua; i kiei aku ka
Kauakahiakahoowaha was therefore startled and looking Waolani, e pii ana ke kahi maha o ke kaua i Pu
around he saw that the enemy was already formed for hoi ko Koolau kaua i Kaniakapupu, a o kekahi
battle. When he looked down the bottom of Waolani, Koolau kaua mai aia ma ka pali o Kalihi, ke pii
one wing of the army was climbing Puuiwa; the army maha o ko Kona aku nei kaua a hookui me ko
from Koolau was coming down Kaniakapupu, while one alaila puni o Kualii.
of the wings of the army from Koolau was already on
the Kalihi cliffs, and still another wing from Kona was
coming up soon to meet the army from Koolau, whereby
Kualii would be entirely surrounded.

When Kauakahiakahoowaha saw this he called out to A ike ae la o Kauakahiakahoowaha i keia mea,
Kualii: “Say, where are you? Are you to continue Kualii: “E, auhea oe? He moe mai ka kau, eia
sleeping, when here we are surrounded by the enemy?” kaua?” A no keia mea, pane ae la o Kualii mai
When Kualii heard this he spoke from within the bed pulou ana: “I aha auanei ka’u pono e ala ae a
clothes that covered his head: “What can I do by getting a’u pono, o ka noho iho no a hiki mai ka luku;
up? There is only one thing for me to do, that is, to kakou make e hele aku ai; oia e, he wahi hapa
remain where I am till the slaughter gets here. What mai i ka olelo, e pono ai ke hele aku i ke kaua
have we on hand to fight them with when we can see no kakou keia kaua.”
for ourselves that they have no end of men on their
side. On the other hand it is entirely within reason that
this battle is not intended for us.”

That morning a messenger was seen coming as though Ia kakahiaka, ike ia aku la kekahi luna i hooun
sent by Lonoikaiaka. He approached Kualii and said: Lonoikaika mai, a hele aku la a halawai me Ku
“There is going to be a battle today.” When Kualii heard aku: “He kaua ko keia la.” A lohe o Kualii i keia
the messenger he replied: “Why did they send you? If alaila i aku la i ka luna: “I hoouna ia mai la no
you wish to make war come and do so, I shall not no hoi paha e makemake no ke kaua, aole a’u
prevent it. You know well enough that I have not as yet ua ike mai la no ia aole wau i a’o i ke kaua; a
acquired the art of warfare. All would have been well if o ke kala ana ia Oahu nei a puni e kii mai ia’u
there was reason for this. With all this lack of reason, no hoi ia ina la he hala kekahi; me ia hala ole
still you come and make war on a mere youngster la no ia e luku i ke kamalii aole i oo ka iwi. E h
whose bones are not even matured. You go back and Lonoikaika i kuu hewa.” I mai o Hema ka luna
ask Lonoikaika what is my fault.” Hema, the messenger, hewa. O ka lilo ana o ka heiau nau e kapu, ao
replied: “I have heard of your fault. It is the fact that heiau ia oe, he akua ka ka mea nana e kapu k
you dedicated the temple, taking upon yourself Kualii: “O hoi a hai aku ia Lonoikaikaole, na’u e
something only a god has the right to do.” Kualii replied:
“Go back and tell Lonoikaikaole 316 that I have the right
to dedicate this temple.”

Hema thereupon returned to the king and reported as Hoi aku la o Hema, a olelo aku la i ke alii: “I i
follows: “Kualii told me to come back and tell you the hoi mai au a olelo aku ia oe, penei oia i olelo m
following words: ‘Go back and tell Lonoikaikaole that I ia Lonoikaikaole, na’u e kapu keia heiau.’ A pe
have the right to dedicate this temple.’ ” When mai nei ia’u.” A lohe o Lonoikaika i keia olelo [4
Lonoikaika heard what Hema had to say, he became manawa, aole o kana mai a ka huhu o Lonoika
very angry and remarked: “Is this youngster who is still “Eia ka oia wahi keiki ma’i lewalewa ko’u mea
so young that he has no knowledge of what shame is, ka ikaika ole. A heaha la hoi kana.”
going to be the one to tell me that I am [412]not strong
enough? Well, we’ll see about it.” Lonoikaika then sent
Hema to hasten and inform the army from Koolau to
bring the wings of the armies together so as to surround
Kualii.

When the armies were ready to begin the conflict, Kualii Hoouna aku la o Lonoikaika ia Hema e holo e
looked about him and saw that the different armies were kaua mai, e hui na holo o na kaua, e hoopuni
closing in on him, and the grass was so thickly covered makaukau ke kaua, ia manawa, nana aku la o
with men that it was dried up from the tramping; he ana na kaua a pau, ua owela ka nahele; alaila
then remarked to his own personal attendant, ia Maheleana i kona kahu ponoi: “E, auhea oe
Maheleana: “Say, where are you? This morning you ao ke kaua, ao ke koa.” I aku o Maheleana: “A
must learn how to fight and how to be brave.” keia pakela nui, ke poai mai nei ka ohu mauka
Maheleana replied: “One cannot show his strength ma o.” I aku o Kualii: “Elua kaua, elua o Kane
against such odds. The rain clouds are encircling from no kakou la, o ka pau no ia o ia nui i ka hee.”
above, from sea-ward and from all sides.” Kualii spoke aku ana ia Kualii, komo ae la oia i ka heiau e p
up: “There are two of us as Kane and Kanaloa are also ka pule ana, i nana aku ka hana o Maheleana,
two. Let us then make a stand and you will see these kaua. Ia manawa, lalau ae la o Kualii i kana la
numbers flee.” While the armies were closing around Manaiakalani, a haawi aku la ia Maheleana, m
Kualii he entered the temple to pray. At the close of laau palau, puka aku oe a komo iloko o ke kau
Kualii’s prayer Maheleana looked and lo, the enemy was
close upon them. Kualii then reached for his war club
Manaiakalani 317 and handed it to Maheleana with the
remark: “Here is my war club, go out and enter into the
army of Lonoikaika.”

As directed by Kualii, Maheleana went forth and began E like me ka Kualii olelo, puka ae la oia a luku
the slaughter of the people with such courage that the palau, a hee aku la ke kaua ma kona alo pono
enemy retired from before him and ran directly toward Lonoikaika; na lakou i hee, hee na kaua a pun
Lonoikaika. When these people withdrew the whole of ka pali o Waolani, hiolo aku la lakou me he iliil
the enemy retreated, those on the pali of Waolani fell luku aku la o Kualii a pau loa na ’lii ma ko Lon
over like pebbles down the pali. Kualii then slew almost aku la na kanaka make me he pauku laau la, k
all the chiefs on Lonoikaika’s side. The dead bodies were make ia kaua ana; a lanakila ae o Kualii ma ia
strewn around like logs of wood, so great was the oia ka hoouka kaua mua a Kualii, a lilo ae la ia
number of those that were killed in this battle. Kualii Moanalua a Maunalua. Mahope iho o ia manaw
was therefore victorious in this his first battle and he Kualii i Kailua ma Koolaupoko, a malaila oia i n
became the owner of all the land from Moanalua to kona hale nui o Kalanihale.
Maunaloa. Shortly after this Kualii went and lived in
Kailua, Koolaupoko, in a great palace called Kalanihale.

Sometime after this, Kualii and Maheleana, his personal I kekahi manawa mahope mai, ao ae la o Kua
attendant and fellow companion in battle, took lessons Maheleana, kona kahu, a hoa kaua pu hoi, i ka
in learning the art of using the war club, and he took a lawe ae la oia ia Kahai, a me Malanaihaehae
Kahai and Malanaihaehae to be his chief warriors. They nona, ao iho la lakou nei i na mea kaua a pau
all studied the different arts of warfare until they were manawe mahope iho, holo ae la o Kualii me ko
quite proficient. Shortly after this Kualii and his chief Kauai, no ka manao e loaa ia lakou ka laau pa
warriors sailed for Kauai, being desirous of procuring malaila. A ma ia hele ana, ua loaa ko lakou ma
certain kinds of war clubs. 318 On this tour they were able la lakou me ua mau laau palau la, a kapa aku
to obtain what they wanted and returned with their new laau o Hulimokualana.
weapons. Kualii named his war club Hulimokualana. 319

On their return from Kauai, Kualii desired to land at Hoi mai la o Kualii mai Kauai mai a pae ma Ka
Kamaile, Waianae, but upon arrival there he found that hoi, ua hoonoho mua ia ke kaua e ko Waianae
the place was already prepared for battle under the o ko Koolauloa alii mai me kona kaua. E kali a
command of the chief of Waianae and Ewa, the Koolau Kualii mai Kauai mai, alaila o ka hoomaka no i
chief and his army had also arrived there and all were
waiting for Kualii’s return from Kauai when they would
engage him.

While out at sea some distance from land Kualii, by his A waena moana, ike e no o Kualii ua puni o W
supernatural powers, knew beforehand that Waianae hoomoemoe. Ia manawa, olelo aku la o Kualii
was surrounded by an army which was waiting for him. laua me Maheleana: “E, ua paa uka o Waianae
So he remarked to Malanaihaehae and Maheleana: “Say, no ia kakou a pae o ka hoomaka no ia o ka luk
Waianae is surrounded by an army that is ready to fight ike e no o Kualii i keia mau mea ma kona ano
us as soon as we make a landing.” Before Kualii had mamua o kona holo ana i Kauai, ua kauoha oi
sailed for Kauai he ordered his men to come and meet e imi mua mai, ke [415]hiki aku i na la e hoi ak
him at Waianae upon his [414]return from Kauai, but Kauai mai. Aka, ia Kualii ma i pae aku ai ma K
when Kualii and his fellow travelers arrived outside of puni i ke kaua; a no ia mea, lana iho la na waa
Kamaile they saw the place surrounded by an army. ia la, a ao ia po. Ia kakahiaka ana ae, i nana a
Upon seeing this they laid off in their canoes all that day Kualii ma, ua uhi ia mai ke pili o Kamaile e na
and night. In the morning when Kualii looked he saw the
pili grass of Kamaile was completely covered by the
people.

While on the canoes that morning Kualii, upon seeing Iluna no o na waa ma ia kakahiaka, pane aku
the people, addressed them in the following words: “You makemake kaua me Kualii, ihea e kaua ai?” O
no doubt want to fight Kualii, but where will the battle “Pae no na waa i uka nei, o ke kaua no ia.” I a
be?” The people from the shore replied: “As soon as the aku kakou iuka o Kalena, ilaila e kaua ai. Ina i
canoes land the fighting will commence.” Kualii ka moana loa no e hele ai na waa, o Molokai k
answered back: “Let us go to Kalena and fight there. If olelo a Kualii, iuka o Kalena e kaua ai, ae aku
you insist on fighting here the canoes will continue by alii. A nolaila, pae aku la lakou iuka, o Kualii, o
sea and land at Molokai.” Because of this request of Malanaihaehae. Hele aku la lakou mauka a no
Kualii to go to Kalena and there fight, the chiefs of po a ao ae, ma ke kakahiakanui, o ka hoouka
Waianae consented because it was but a reasonable kaua ma Kalena, i ke kula o Haleauau ma Lihu
request. Kualii, Maheleana and Malanaihaehae therefore he ekolu mano ka nui; a ma ko Kualii aoao ho
came ashore and proceeded by land to Malamanui. All lakou, a o ka hee iho la no ia o na alii o Waian
that night both sides took a long rest; but early in the a kapa iho la o Kualii ia hoouka kaua ana o Ka
morning the fighting commenced at Kalena on the plains
of Haleauau, at Lihue. On the one side there were
twelve thousand men, while on Kualii’s side there were
but three men, and yet the armies of the chiefs of
Waianae and Koolauloa were routed. Kualii named this
the battle of Kalena.

A few days after this three more battles were fought, at A ma kekahi mau la ae, he mau hoouka kaua
Malamanui, Pulee and Paupauwela. These were the Pulee, a ma Paupauwela, oia ka hoouka nui an
greatest of the battles fought by Kualii in all the Oahu mamua o na hoouka kaua a pau ma Oahu nei
contests. Sometime after he had conquered the whole manawa mahope mai o kona ai ana ia Oahu, l
of Oahu he heard that there was a battle in Hilo, Hawaii; kaua ma Hilo i Hawaii; nolaila, hele aku la oia
he therefore made up his mind to make a trip to Hawaii mau pukaua. A lohe o Haalilo, ua hiki aku o Ku
with his chief warriors. When Haalilo heard that Kualii Laupahoehoe, alaila, hoomakaukau ae la oia n
had arrived at Laupahoehoe he immediately prepared aku la o Kualii i Peahi ma Hilo, halawai koke ae
for war, so that when Kualii reached Peahi in Hilo he ran Haalilo, a o ka hoomaka iho la no ia o ke kaua
into Haalilo and the battle commenced. It was of but Kualii; a lohe aku la na ’lii o Puna, ua hee o Ha
short duration and Kualii was victorious. When the chiefs aku la ko Puna mau alii.
of Puna heard that Haalilo was beaten they too fell back.

Shortly after this word was brought to Kualii at Hilo that A mahope iho o ia manawa, hiki aku ka lohe ia
the chiefs of Oahu had again risen against him and were na ’lii o Oahu nei. Ia manawa, hoi mai la o Kua
ready to dispute his title as king of Oahu. Upon hearing hiki ma Oahu nei, ua pau loa na kanaka a pau
this Kualii returned from Hilo to Oahu and found upon Waianae kahi i noho ai, ua hui ae la na ’lii mua
his arrival that all the people, together with the me ka manao lokahi, e kipi aku ia Kualii. A no
rebellious chiefs, had gone to Waianae to hold a council Kualii, aia na ’lii a pau malalo o Waianae; nola
of war with the one set purpose of fighting him. When oia me kona mau pukaua, a hiki lakou malaila,
Kualii heard that all the chiefs were gathered at ka hana, he mea e ka nui o na kanaka. A o ka
Waianae, he continued on with his chief warriors for that no ia, a lanakila iho la o Kualii ma ia hoouka a
place. Upon arrival at the seat of war they looked and aku a Kualii ma i ka nui i make ma ia hoouka a
saw that the rebellious chiefs had indeed a very large wai o Kalapo, a nui no hoi na kupapau malalo
army. No time was lost, however, for the battle hoike ia kekahi mau lalani mele ma ka haku an
immediately commenced, and again Kualii was e like me ke ano o ia hoouka kaua ana malaila
victorious. After the battle Kualii and his chief warriors
looked over the battle ground and saw that a very large
number of men had been killed, so much so that the
waters of Kalapo were dammed and a large number of
dead bodies were strewn below Eleu. Because of this
great victory certain lines of mele were composed by his
attendants which read as follows:

A battle for Ku, He kaua na Ku,

Beating his enemy on the heights of Kawaluna. E uhau ana iluna o Kawaluna
Where, where is the battle field Ihea, ihea la ke kahua,
Where the warrior is to fight? Paio ai o ke koa-a?
On the field of Kalena, 5 I ka i kahua i Kalena, 5
At Manini, at Hanini, I Manini, i Hanini
Where was poured the water of the god I ninia i ka wai akua,
At Kahana, at Malamanui; I Kahana i Malamanui
On the heights of Kapapa, at Paupauwela, Ka luna o Kapapa, i Paupauw
Where they lean and rest; 10 I ka hilinai i ke kalele, 10
At the hala trees of indolent Halahalanui, [416] Ka hala o Halahalanui maaue
At the ohia grove of Pule-e, Ke kula ohia ke Pule-e, [417]
The god of Lono, of Makalii, Ke ’kua o Lono o Makalii
The fragrant branch of the Ukulonoku, Ka lala aalao Ukulonoku,
Mayhap from Kona, from Lihue, 15 No Kona paha, no Lihue. 15
For the day at Maunauna, No ka la i Maunauna,
For the water at Paupauwela; No ka wai i Paupauwela,
Growing low at Nepee, I ulu haa lilo i Nepee,
At the slaughter of Aui, A ka hauna o Aui.
Where the priests joined in the battle. 20 Kokomo kahuna i kakua laau,
Ku is arrayed in his feather cloak, Komo Ku i kona ahuula,
The sun-lighted rain in the heavens, Ka wela o ka ua i ka lani,
The sun at Kauakahihale. Ka la i Kauakahihale,
Red is the leaf of the mamane, Ula ka lau o ka mamane,
The koaie of Kauai; 25 Ke koaie o Kauai; 25
The sea grass has been stripped by Ku— He pili kai ihi ia e Ku,
The waving [grass] of Kamaile; Ka aloalo o Kamaile,
The towering surf of Maihiwa, Ka nalu kakala o Maihiwa,
Which dammed up the waters of Halapo. Pania ka wai i Halapo,
The breaking up is below at Eleu, 30 Ka naha ilalo o Eleu. 30
The rain is drawn away to the sky, Huki kaua a moa i ka lani,
Like a full retreat from the mountain; Me he hee nui no kuahiwi;
It must be the defeat of Hilo by Puna, Ka hee ana o Hilo ia Puna,
There at Hilo is Peahi. Aia ma Hilo Peahi;
Red is the water of Paupauwela, 35 Ula ka wai i Paupauwela, 35
From the slain at Malamanui, Ke kilau o Malamanui,
The slain on the ridge at Kapapa. Ka moo kilau i Kapapa.
The tidings reached Haalilo Kui ka lono ia Haalilo,
Your younger brother is beaten. Kaua aku la ko kaina;
Haalilo is sore at heart, 40 Hahai Haalilo i ka manawa; 4
For Ku has left but few of the priests; I kai muku kahuna ia Ku;
They are beaten by Ku, I la ka mawa ia Ku,
The children of Haalilo. I keiki a Haalilo.
Here is Malanaihaehae, Eia Malanaihaehae,
Offspring of mischief-making Niheu, 45 Kama a Niheu kalohe, 45
The dammer of the waters of Kekeuna. Ke pani wai o Kekuna,
A prodigy among the people. He mee nei no ke kanaka,
He is girding on his robe, Ke pu nei i ka aahu,
He is whirling his weapon [in the air], Ke lapa nei i ka laau,
The war club is caught in his robe. 50 Ka laulau o ka aahu, 50
Here is Haalilo, Eia Haalilo-e!—
Ku is indeed king. O Ku no ke alii.