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COVID-19 infection: Origin,

transmission, and characteristics of


human coronaviruses Muhammad
Adnan Shereen
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Journal of Advanced Research 24 (2020) 91–98

Contents lists available at ScienceDirect

Journal of Advanced Research


journal homepage: www.elsevier.com/locate/jare

COVID-19 infection: Origin, transmission, and characteristics of human


coronaviruses
Muhammad Adnan Shereen a,b,1, Suliman Khan a,1,⇑, Abeer Kazmi c, Nadia Bashir a, Rabeea Siddique a
a
The Department of Cerebrovascular Diseases, The Second Affiliated Hospital of Zhengzhou University, Zhengzhou, PR China
b
State Key Laboratory of Virology, College of Life Sciences, Wuhan University, Wuhan, PR China
c
College of Life Sciences, Wuhan University, Wuhan, PR China

g r a p h i c a l a b s t r a c t

a r t i c l e i n f o a b s t r a c t

Article history: The coronavirus disease 19 (COVID-19) is a highly transmittable and pathogenic viral infection caused by
Received 15 March 2020 severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), which emerged in Wuhan, China and
Accepted 15 March 2020 spread around the world. Genomic analysis revealed that SARS-CoV-2 is phylogenetically related to sev-
Available online 16 March 2020
ere acute respiratory syndrome-like (SARS-like) bat viruses, therefore bats could be the possible primary
reservoir. The intermediate source of origin and transfer to humans is not known, however, the rapid
Keywords: human to human transfer has been confirmed widely. There is no clinically approved antiviral drug or
Coronaviruses
vaccine available to be used against COVID-19. However, few broad-spectrum antiviral drugs have been
COVID-19
Origin
evaluated against COVID-19 in clinical trials, resulted in clinical recovery. In the current review, we sum-
Outbreak marize and comparatively analyze the emergence and pathogenicity of COVID-19 infection and previous
Spread human coronaviruses severe acute respiratory syndrome coronavirus (SARS-CoV) and middle east respi-
ratory syndrome coronavirus (MERS-CoV). We also discuss the approaches for developing effective vac-
cines and therapeutic combinations to cope with this viral outbreak.
Ó 2020 THE AUTHORS. Published by Elsevier BV on behalf of Cairo University. This is an open access article
under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Introduction
Peer review under responsibility of Cairo University.
⇑ Corresponding author. Coronaviruses belong to the Coronaviridae family in the Nidovi-
E-mail address: suliman.khan18@mails.ucas.ac.cn (S. Khan). rales order. Corona represents crown-like spikes on the outer sur-
1
Contributed equally (M.A.S and S.K). face of the virus; thus, it was named as a coronavirus.

https://doi.org/10.1016/j.jare.2020.03.005
2090-1232/Ó 2020 THE AUTHORS. Published by Elsevier BV on behalf of Cairo University.
This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
92 M.A. Shereen et al. / Journal of Advanced Research 24 (2020) 91–98

Coronaviruses are minute in size (65–125 nm in diameter) and nationals were diagnosed to be infected with another coronavirus.
contain a single-stranded RNA as a nucleic material, size ranging The detected virus was confirmed as a member of coronaviruses
from 26 to 32kbs in length (Fig. 1). The subgroups of coronaviruses and named as the Middle East Respiratory Syndrome Coronavirus
family are alpha (a), beta (b), gamma (c) and delta (d) coronavirus. (MERS-CoV). The World health organization reported that MERS-
The severe acute respiratory syndrome coronavirus (SARS-CoV), coronavirus infected more than 2428 individuals and 838 deaths
H5N1 influenza A, H1N1 2009 and Middle East respiratory syn- [8]. MERS-CoV is a member beta-coronavirus subgroup and phylo-
drome coronavirus (MERS-CoV) cause acute lung injury (ALI) and genetically diverse from other human-CoV. The infection of MERS-
acute respiratory distress syndrome (ARDS) which leads to pul- CoV initiates from a mild upper respiratory injury while progres-
monary failure and result in fatality. These viruses were thought sion leads to severe respiratory disease. Similar to SARS-
to infect only animals until the world witnessed a severe acute res- coronavirus, patients infected with MERS-coronavirus suffer pneu-
piratory syndrome (SARS) outbreak caused by SARS-CoV, 2002 in monia, followed by ARDS and renal failure [9].
Guangdong, China [1]. Only a decade later, another pathogenic Recently, by the end of 2019, WHO was informed by the Chi-
coronavirus, known as Middle East respiratory syndrome coron- nese government about several cases of pneumonia with unfamil-
avirus (MERS-CoV) caused an endemic in Middle Eastern countries iar etiology. The outbreak was initiated from the Hunan seafood
[2]. market in Wuhan city of China and rapidly infected more than
Recently at the end of 2019, Wuhan an emerging business hub 50 peoples. The live animals are frequently sold at the Hunan sea-
of China experienced an outbreak of a novel coronavirus that killed food market such as bats, frogs, snakes, birds, marmots and rabbits
more than eighteen hundred and infected over seventy thousand [10]. On 12 January 2020, the National Health Commission of China
individuals within the first fifty days of the epidemic. This virus released further details about the epidemic, suggested viral pneu-
was reported to be a member of the b group of coronaviruses. monia [10]. From the sequence-based analysis of isolates from the
The novel virus was named as Wuhan coronavirus or 2019 novel patients, the virus was identified as a novel coronavirus. Moreover,
coronavirus (2019-nCov) by the Chinese researchers. The Interna- the genetic sequence was also provided for the diagnosis of viral
tional Committee on Taxonomy of Viruses (ICTV) named the virus infection. Initially, it was suggested that the patients infected with
as SARS-CoV-2 and the disease as COVID-19 [3–5]. In the history, Wuhan coronavirus induced pneumonia in China may have visited
SRAS-CoV (2003) infected 8098 individuals with mortality rate of the seafood market where live animals were sold or may have used
9%, across 26 contries in the world, on the other hand, novel corona infected animals or birds as a source of food. However, further
virus (2019) infected 120,000 induviduals with mortality rate of investigations revealed that some individuals contracted the infec-
2.9%, across 109 countries, till date of this writing. It shows that tion even with no record of visiting the seafood market. These
the transmission rate of SARS-CoV-2 is higher than SRAS-CoV and observations indicated a human to the human spreading capability
the reason could be genetic recombination event at S protein in the of this virus, which was subsequently reported in more than 100
RBD region of SARS-CoV-2 may have enhanced its transmission countries in the world. The human to the human spreading of
ability. In this review article, we discuss the origination of human the virus occurs due to close contact with an infected person,
coronaviruses briefly. We further discuss the associated infectious- exposed to coughing, sneezing, respiratory droplets or aerosols.
ness and biological features of SARS and MERS with a special focus These aerosols can penetrate the human body (lungs) via inhala-
on COVID-19. tion through the nose or mouth (Fig. 2) [11–14].

Comparative analysis of emergence and spreading of


coronaviruses Primary reservoirs and hosts of coronaviruses

In 2003, the Chinese population was infected with a virus caus- The source of origination and transmission are important to be
ing Severe Acute Respiratory Syndrome (SARS) in Guangdong pro- determined in order to develop preventive strategies to contain the
vince. The virus was confirmed as a member of the Beta- infection. In the case of SARS-CoV, the researchers initially focused
coronavirus subgroup and was named SARS-CoV [6,7]. The infected on raccoon dogs and palm civets as a key reservoir of infection.
patients exhibited pneumonia symptoms with a diffused alveolar However, only the samples isolated from the civets at the food
injury which lead to acute respiratory distress syndrome (ARDS). market showed positive results for viral RNA detection, suggesting
SARS initially emerged in Guangdong, China and then spread that the civet palm might be secondary hosts [15]. In 2001 the
rapidly around the globe with more than 8000 infected persons samples were isolated from the healthy persons of Hongkong and
and 776 deceases. A decade later in 2012, a couple of Saudi Arabian the molecular assessment showed 2.5% frequency rate of anti-
bodies against SARS-coronavirus. These indications suggested that
SARS-coronavirus may be circulating in humans before causing the
outbreak in 2003 [16]. Later on, Rhinolophus bats were also found
to have anti-SARS-CoV antibodies suggesting the bats as a source of
viral replication [17]. The Middle East respiratory syndrome
(MERS) coronavirus first emerged in 2012 in Saudi Arabia [9].
MERS-coronavirus also pertains to beta-coronavirus and having
camels as a zoonotic source or primary host [18]. In a recent study,
MERS-coronavirus was also detected in Pipistrellus and Perimyotis
bats [19], proffering that bats are the key host and transmitting
medium of the virus [20,21]. Initially, a group of researchers sug-
gested snakes be the possible host, however, after genomic similar-
ity findings of novel coronavirus with SARS-like bat viruses
supported the statement that not snakes but only bats could be
the key reservoirs (Table 1) [22,23]. Further analysis of homolo-
gous recombination revealed that receptor binding spike glycopro-
tein of novel coronavirus is developed from a SARS-CoV (CoVZXC21
Fig. 1. Structure of respiratory syndrome causing human coronavirus. or CoVZC45) and a yet unknown Beta-CoV [24]. Nonetheless, to
M.A. Shereen et al. / Journal of Advanced Research 24 (2020) 91–98 93

Fig. 2. The key reservoirs and mode of transmission of coronaviruses (suspected reservoirs of SARS-CoV-2 are red encircled); only a and b coronaviruses have the ability to
infect humans, the consumption of infected animal as a source of food is the major cause of animal to human transmission of the virus and due to close contact with an
infected person, the virus is further transmitted to healthy persons. Dotted black arrow shows the possibility of viral transfer from bat whereas the solid black arrow
represent the confirmed transfer. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

Table 1
Comparative analysis of biological features of SARS-CoV and SARS-CoV-2.

Features SARS-CoV SARS-CoV-2 Reference


Emergence date November 2002 December 2019 [37,79–81]
Area of emergence Guangdong, China Wuhan, China
Date of fully controlled July 2003 Not controlled yet
Key hosts Bat, palm civets and Raccon dogs Bat [22,82,83]
Number of countries infected 26 109 [84]
Entry receptor in humans ACE2 receptor ACE2 receptor [22,55,85]
Sign and symptoms fever, malaise, myalgia, headache, diarrhoea, shivering, Cough, fever and shortness of breath [12,23,85]
cough and shortness of breath
Disease caused SARS, ARDS SARS, COVID-19 [85,86]
Total infected patients 8098 123882 [84]
Total recovered patients 7322 67051
Total died patients 776 (9.6% mortality rate) 4473 (3.61% mortality rate)

eradicate the virus, more work is required to be done in the aspects teins, and membrane proteins, such as RNA polymerase, 3-
of the identification of the intermediate zoonotic source that chymotrypsin-like protease, papain-like protease, helicase, glyco-
caused the transmission of the virus to humans. protein, and accessory proteins [30,31]. The spike protein of
SARS-CoV-2 contains a 3-D structure in the RBD region to maintain
the van der Waals forces [32]. The 394 glutamine residue in the
Key features and entry mechanism of human coronaviruses RBD region of SARS-CoV-2 is recognized by the critical lysine 31
residue on the human ACE2 receptor [33]. The entire mechanism
All coronaviruses contain specific genes in ORF1 downstream of pathogenicity of SARS-CoV-2, from attachment to replication is
regions that encode proteins for viral replication, nucleocapsid well mentioned in Fig. 3.
and spikes formation [25]. The glycoprotein spikes on the outer
surface of coronaviruses are responsible for the attachment and
entry of the virus to host cells (Fig. 1). The receptor-binding Genomic variations in SARS-CoV-2
domain (RBD) is loosely attached among virus, therefore, the virus
may infect multiple hosts [26,27]. Other coronaviruses mostly rec- The genome of the SARS-CoV-2 has been reported over 80%
ognize aminopeptidases or carbohydrates as a key receptor for identical to the previous human coronavirus (SARS-like bat CoV)
entry to human cells while SARS-CoV and MERS-CoV recognize [34]. The Structural proteins are encoded by the four structural
exopeptidases [2]. The entry mechanism of a coronavirus depends genes, including spike (S), envelope (E), membrane (M) and nucle-
upon cellular proteases which include, human airway trypsin-like ocapsid (N) genes. The orf1ab is the largest gene in SARS-CoV-2
protease (HAT), cathepsins and transmembrane protease serine 2 which encodes the pp1ab protein and 15 nsps. The orf1a gene
(TMPRSS2) that split the spike protein and establish further pene- encodes for pp1a protein which also contains 10 nsps [34–36].
tration changes [28,29]. MERS-coronavirus employs dipeptidyl According to the evolutionary tree, SARS-CoV-2 lies close to the
peptidase 4 (DPP4), while HCoV-NL63 and SARS-coronavirus group of SARS-coronaviruses [37,38] (Fig. 5). Recent studies have
require angiotensin-converting enzyme 2 (ACE2) as a key receptor indicated notable variations in SARS-CoV and SARS-CoV-2 such
[2,26]. as the absence of 8a protein and fluctuation in the number of
SARS-CoV-2 possesses the typical coronavirus structure with amino acids in 8b and 3c protein in SARS-CoV-2 [34] (Fig. 4). It is
spike protein and also expressed other polyproteins, nucleopro- also reported that Spike glycoprotein of the Wuhan coronavirus
94 M.A. Shereen et al. / Journal of Advanced Research 24 (2020) 91–98

Fig. 3. The life cycle of SARS-CoV-2 in host cells; begins its life cycle when S protein binds to the cellular receptor ACE2. After receptor binding, the conformation change in
the S protein facilitates viral envelope fusion with the cell membrane through the endosomal pathway. Then SARS-CoV-2 releases RNA into the host cell. Genome RNA is
translated into viral replicase polyproteins pp1a and 1ab, which are then cleaved into small products by viral proteinases. The polymerase produces a series of subgenomic
mRNAs by discontinuous transcription and finally translated into relevant viral proteins. Viral proteins and genome RNA are subsequently assembled into virions in the ER
and Golgi and then transported via vesicles and released out of the cell. ACE2, angiotensin-converting enzyme 2; ER, endoplasmic reticulum; ERGIC, ER–Golgi intermediate
compartment.

is modified via homologous recombination. The spike glycoprotein another suitable option to develop spike glycoprotein targeting
of SARS-CoV-2 is the mixture of bat SARS-CoV and a not known therapeutics against novel coronaviruses. Recently, mice models
Beta-CoV [38]. In a fluorescent study, it was confirmed that the and clinical isolates were used to develop any therapeutic strategy
SARS-CoV-2 also uses the same ACE2 (angiotensin-converting against SARS-CoV-2 induced COVID-19 [46,47]. In a similar study,
enzyme 2) cell receptor and mechanism for the entry to host cell artificial intelligence prediction was used to investigate the inhibi-
which is previously used by the SARS-CoV [39,40]. The single tory role of the drug against SARS-CoV-2 [48]. SARS-CoV-2 infected
N501T mutation in SARS-CoV-2’s Spike protein may have signifi- patients were also used to conduct randomized clinical trials
cantly enhanced its binding affinity for ACE2 [33]. [46,49,50]. It is now important that the scientists worldwide col-
laborate the design a suitable model and investigate the in vivo
The major obstacle in research progress mechanisms associated with pathogenesis of SARS-CoV-2.

Animal models play a vital role to uncover the mechanisms of Potential therapeutic strategies against COVID-19
viral pathogenicity from the entrance to the transmission and
designing therapeutic strategies. Previously, to examine the repli- Initially, interferons-a nebulization, broad-spectrum antibi-
cation of SARS-CoV, various animal models were used which otics, and anti-viral drugs were used to reduce the viral load
showed the symptoms of severe infection [43]. In contrast to [49,51,52], however, only remdesivir has shown promising impact
SARS-CoV, no MERS-CoV pathogenesis was observed in small ani- against the virus [53]. Remdesivir only and in combination with
mals. Mice are not vulnerable to infection by MERS-coronavirus chloroquine or interferon beta significantly blocked the SARS-
due to the non-compatibility of the DPP4 receptor [44]. As the CoV-2 replication and patients were declared as clinically recov-
entire genome of the 2019-novel coronavirus is more than 80% ered [46,50,52]. Various other anti-virals are currently being eval-
similar to the previous human SARS-like bat CoV, previously used uated against infection. Nafamostat, Nitazoxanide, Ribavirin,
animal models for SARS-CoV can be utilized to study the infectious Penciclovir, Favipiravir, Ritonavir, AAK1, Baricitinib, and Arbidol
pathogenicity of SARS-CoV-2. The human ACE2 cell receptor is rec- exhibited moderate results when tested against infection in
ognized by both SARS and Novel coronaviruses. Conclusively, patients and in-vitro clinical isolates [46,48,50,52]. Several other
TALEN or CRISPR-mediated genetically modified hamsters or other combinations, such as combining the antiviral or antibiotics with
small animals can be utilized for the study of the pathogenicity of traditional Chinese medicines were also evaluated against SARS-
novel coronaviruses. SARS-CoV has been reported to replicate and CoV-2 induced infection in humans and mice [46]. Recently in
cause severe disease in Rats (F344), where the sequence analysis Shanghai, doctors isolated the blood plasma from clinically recov-
revealed a mutation at spike glycoprotein [45]. Thus, it could be ered patients of COVID-19 and injected it in the infected patients
M.A. Shereen et al. / Journal of Advanced Research 24 (2020) 91–98 95

Fig. 4. Betacoronaviruses genome organization; The Betacoronavirus for human (SARS-CoV-2, SARS-CoV and MERS-CoV) genome comprises of the 50 -untranslated region (50 -
UTR), open reading frame (orf) 1a/b (green box) encoding non-structural proteins (nsp) for replication, structural proteins including spike (blue box), envelop (maroon box),
membrane (pink box), and nucleocapsid (cyan box) proteins, accessory proteins (light gray boxes) such as orf 3, 6, 7a, 7b, 8 and 9b in the SARS-CoV-2 genome, and the 30 -
untranslated region (30 -UTR). The doted underlined in red are the protein which shows key variation between SARS-CoV-2 and SARS-CoV. The length of nsps and orfs are not
drawn in scale. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

Fig. 5. Phylogenetic tree of coronaviruses (content in red is the latest addition of newly emerged SARS-CoV-2 and WSFMP Wuhan-Hu-1 is used as a reference in the tree); The
phylogenetic tree showing the relationship of Wuhan-Hu-1 (denoted as red) to selected coronavirus is based on nucleotide sequences of the complete genome. The viruses
are grouped into four genera (prototype shown): Alphacoronavirus (sky blue), Betacoronavirus (pink), Gammacoronavirus (green) and Deltacoronavirus (light blue).
Subgroup clusters are labeled as 1a and 1b for the Alphacoronavirus and 2a, 2b, 2c, and 2d for the Betacoronavirus. This tree is based on the published trees of Coronavirinae
[3,41] and reconstructed with sequences of the complete RNA- dependent RNA polymerase- coding region of the representative novel coronaviruses (maximum likelihood
method using MEGA 7.2 software). severe acute respiratory syndrome coronavirus (SARS- CoV); SARS- related coronavirus (SARSr- CoV); the Middle East respiratory
syndrome coronavirus (MERS- CoV); porcine enteric diarrhea virus (PEDV); Wuhan seafood market pneumonia (Wuhan-Hu-1). Bat CoV RaTG13 Showed high sequence
identity to SARS-CoV-2 [42]. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

who showed positive results with rapid recovery [54]. In a recent receptor-binding motif. CR3022 has the potential to be developed
study, it was identified that monoclonal antibody (CR3022) binds - as a therapeutic candidate, alone or in combination with other
with the spike RBD of SARS-CoV-2. This is likely due to the anti- neutralizing antibodies for the prevention and treatment of -
body’s epitope not overlapping with the divergent ACE2 COVID-19 infection [55].
96 M.A. Shereen et al. / Journal of Advanced Research 24 (2020) 91–98

Vaccines for SARS-CoV-2 Most importantly, human coronaviruses targeting vaccines and
antiviral drugs should be designed that could be used against the
There is no available vaccine against COVID-19, while previous current as well as future epidemics. There are many companies
vaccines or strategies used to develop a vaccine against SARS-CoV working for the development of effective SARS-CoV-2 vaccines,
can be effective. Recombinant protein from the Urbani (AY278741) such as Moderna Therapeutics, Inovio Pharmaceuticals, Novavax,
strain of SARS-CoV was administered to mice and hamsters, Vir Biotechnology, Stermirna Therapeutics, Johnson & Johnson,
resulted in the production of neutralizing antibodies and protec- VIDO-InterVac, GeoVax-BravoVax, Clover Biopharmaceuticals, Cur-
tion against SARS-CoV [56,57]. The DNA fragment, inactivated eVac, and Codagenix. But there is a need for rapid human and
whole virus or live-vectored strain of SARS-CoV (AY278741), sig- animal-based trails as these vaccines still require 3–10 months
nificantly reduced the viral infection in various animal models for commercialization. There must be a complete ban on utilizing
[58–63]. Different other strains of SARS-CoV were also used to pro- wild animals and birds as a source of food. Beside the development
duce inactivated or live-vectored vaccines which efficiently of most efficient drug, a strategy to rapidly diagnose SARS-CoV-2 in
reduced the viral load in animal models. These strains include, suspected patient is also required. The signs and symptoms of
Tor2 (AY274119) [64,65], Utah (AY714217) [66], FRA (AY310120) SARS-CoV-2 induced COVID-19 are a bit similar to influenza and
[59], HKU-39849 (AY278491) [57,67], BJ01 (AY278488) [68,69], seasonal allergies (pollen allergies). Person suffering from influ-
NS1 (AY508724) [70], ZJ01 (AY297028) [70], GD01 (AY278489) enza or seasonal allergy may also exhibit temprature which can
[69] and GZ50 (AY304495) [71]. However, there are few vaccines be detected by thermo-scanners, hence the person will become
in the pipeline against SARS-CoV-2. The mRNA based vaccine pre- suspected. Therefore, an accurate and rapid diagnostic kit or meter
pared by the US National Institute of Allergy and Infectious Dis- for detection of SARS-CoV-2 in suspected patients is required, as
eases against SARS-CoV-2 is under phase 1 trial [72]. INO-4800- the PCR based testing is expensive and time consuming. Different
DNA based vaccine will be soon available for human testing [73]. teams of Chinese doctors should immediately sent to Eurpean
Chinese Centre for Disease Control and Prevention (CDC) working and other countries, especially spain and Italy to control the over
on the development of an inactivated virus vaccine [74,75]. Soon spread of COVID-19, because Chinese doctors have efficiently con-
mRNA based vaccine’s sample (prepared by Stermirna Therapeu- trolled the outbreak in china and limited the mortality rate to less
tics) will be available [76]. GeoVax-BravoVax is working to develop than 3% only. The therapeutic strategies used by Chinese, should
a Modified Vaccina Ankara (MVA) based vaccine [77]. While Clover also be followed by other countries.
Biopharmaceuticals is developing a recombinant 2019-nCoV S pro-
tein subunit-trimer based vaccine [78].
Acknowledgments
Although research teams all over the world are working to
investigate the key features, pathogenesis and treatment options,
The authors acknowledge the Postdoctoral grant from The Second
it is deemed necessary to focus on competitive therapeutic options
Affiliated Hospital of Zhengzhou University (for S.K).
and cross-resistance of other vaccines. For instance, there is a pos-
sibility that vaccines for other diseases such as rubella or measles
can create cross-resistance for SARS-CoV-2. This statement of
Declaration of Competing Interest
cross-resistance is based on the observations that children in china
were found less vulnerable to infection as compared to the elder
The authors of this manuscript declare no conflict of interest.
population, while children are being largely vaccinated for measles
in China.
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Keaukanai, 2, 8.

Keauleinakahi ordered to pierce the double canoe of Kaumaielieli and kill Kana
and Niheu, 444.
sword-fish of Kapepeekauila, 444.
warrior in charge of the ocean, 444.
warrior of Kapepeekauila meets and attacks the double canoe; is struck and
killed by Niheu, 444.

Keaumiki and Keauka, tides or demigods, 160–62.


from Kauai, return with Makolea, 510.
gods of tides, 510.
guardians and attendants on Kaikipaananea, 510.

Keawe, 25, 240, 388, 405;


dwelt at Piilani’s, 240.
Hauoa of, 242.
Lono sacred chief by, 356.
lord of Hawaii, 394.
was given birth, 356.

Keawekekahialiiokamoku, 364.
Kualii likened to, 388, 392.
[xxiv]ruled Hawaii four generations before Kamehameha, 388.
turned salt water into fresh, 388.

Keawenuiaumi, 25, 220, 228, 256, 405.


after the death of, 256.
and party proceed to koa forest, 462.
at Kaipalaoa at Mainele’s arrival, 460.
awards his daughter and land to Pikoiakaalala, 462.
battle of Puumaneo, a rebellion against, 314.
bones of six rebellious chiefs killed at battle of, on Puumaneo, 314–20.
circuits Hawaii with Lonoikamakahiki, 264.
contends with his son for the retention of weapons, etc., 260.
father of Lonoikamakahiki, 256.
god of; in charge of Lono; worshipped by Hauna and Loli; thought to be Kaili,
292.
hears report of son’s orders that his war and game implements be destroyed,
260.
high priest with long hair to below his waist in presence of, 264.
Kaikilani became ruler of Hawaii at death of, 266.
king of Hawaii, cautioned by birds against cutting a hollow tree, 458.
orders four men to carry the basket and builds a house for the god and
Kauakahi, 460–62.
ponders upon the future of his boy, 260.
requests Lono to take charge of government, but he declines, 266.
sends messengers for Mainele, vowing to give him daughter and land if he kill
the birds, 458.
sought out Lonoikamakahiki, 260.
told again by the birds of the hollow tree, 462;
vexed, seeks skillful archers to kill them, 458.
vacancy left by, 270.

Keawewai, Hoamakeikekula enveloped in thick fog, arrives at, 536.


Kalamaula lizard king of, 534.

Keawewaihe, 396.

Keeaumoku, 25, 405.

Keehi, sea for mullet is at, 378.

Keelekoha, 382.

Keeumoku, 25. (See Keeaumoku.)

Kehoni, priest, permitted to save himself, 568.

Keiki a kaua, our son, an elastic term, 500.

Keiki, hookama, 182.

Keinohoomanawanui again sees an armed company and says “Our death is close
upon us,” 466.
credited by Kakuhihewa as the cause of victories, 468.
definition of, 466.
discredited by a farmer for the victories, 468.
fears at dagger sign of being discovered, 466.
fears for the result of Kalelealuaka’s wish, 464.
gains victory in battles with Pueonui’s men, 468.
made an officer of Kakuhihewa’s, 468.
seeing an armed company approaching, fears death, 466.
termed by Kalelealuaka a coward, 466.

Keaka, chiefess; has produced eight; sacred bud of, 240.

Kekaa, 284, 302; fleet of war canoes at, 424.

Kekaha battle at Kalamaula, prepared for, by, 418.


chiefs of Koolau and Kona battle against those of, 418.
chiefs of, value their fishing grounds, 416.
Koi and companions land at; thence to Makaeo, 234.
Koolau (Molokai) chiefs desire, 416.
Paepae, a chief of, visits Maui seeking aid of Kauhi, 416.
rain comes by way of, 396.
section of country from Kawela to Maamomi, 416.

Kekaihawewe, son of Moikeha, 118, 132–34, 144–46.

Kekamakahinuiaiku, 32, 48, 80;


almost dead of hunger, 50;
has a double portion, 98.
of bad temper, thrown into the sea, 36.
pretends friendliness, 38.
throws Aukele into pit of Kamooinanea, 38.

Kekamaluahaku, 24.

Kekauilani, 24, 404.

Kekea, or Albino, 8.

Kekea Kapu, 4.

Kekela, 25, 405.

Kekele, a handsome woman, becomes wife of Kaulu, 532.


hala at, planted for her, 530.

Kekohi, the priest, deserts Hakau, 16.

Kekuaokalani’s stick, 96.

Kekuapoiwa, 25, 405.


Kekuapololi, 396.

Kekuawalu, 394.

Kekupuaiawaawa, 284, 302.

Kekuuna, the waters of, 386, 416.

Keliiokaloa, 25, 220, 228, 405.

Kemamo, above Waipio, 200.

Kemau, 192.

Keohe, 344.

Keohokalani, 370, 404.

Keolewa, a mountain of Kauai, spread low is, 372.


Kiki and party are at, 372.

Keoloewa, Kauai chief victorious in revolt, 152.


Kila returned with, to Kauai, 152.
king Puuonale living at, 538.
prime minister of Kila’s, 152.
sails for Waipio: urges Kila to become King, 152.
spirit chief on Maui, 476.

Keoneoio (Maui), Koi returning from Kauai stayed over at, 232.

Keopu, cave of Umi in, at Kailua, 232.

Keopuolono, Kapolei daughter of, sent to entertain Kualii, 418.

Keoua, chiefs rebelled during reign of, 362.

Kepakailiula, adopted through fear by Kakuhihewa; brought to and given Oahu,


510.
acceding to messengers’ overtures, sails for Oahu with his two wives, landing at
Waikiki, 510.
Aiakoake and Kuaikalolo, elders of, 498.
and Makolea became husband and wife, 502.
and wife do nothing but sleep, 502.
at rush of chiefs and warriors of Maui, drops his war club for a hand encounter,
508.
became the fire that lighted Paliuli, 500.
befriends the king’s crier, Kukaea, 512.
besmears the mat of Kakaalaneo, 504.
boards a canoe and sails for Hana, Maui, 504.
born in Keaau, Puna, as an egg, 498.
by aid of young wife, Kukuipahu furnishes canoes and men for invasion of Maui,
506.
called first-born of the beloved one of, 500.
definition of, 498.
favored by king of Kohala, 504.
friend and Kukaea slay in great numbers and force the people to flee, 514–16.
given the daughter of Kukuipahu for wife, 504.
gives charge of Kauai to his friend, 516.
gives his people choice to remain or return, 508.
gives Makolea’s attendant to foster-parents for wife, 502.
hears of king’s sports and joins therein; in contest, defeats Kaikipaananea, 512.
informs foster-parents and moves to Kohala, 502–04.
in hiding, takes Makolea from Kakaalaneo, 504.
in return for kindness, is given answers to king’s riddles, 512–14.
in small canoe, arrives at Waimea, Kauai, 512.
is stayed in his slaughter by his young wife in arms of her father, 508.
[xxv]landing at Hana, the crowd shout in admiration, 506.
led the fight with uprooted trees and rocks, 508.
left asleep at wife’s departure, 502.
legend of, 498.
makes his foster-fathers become kings of Oahu, 510.
makes Kukuipahu king of Maui, 510.
Makolea, wife of, in surf-riding at Waikiki, is taken to Kauai by Keaumiki and
Keauka, 510.
meets and is befriended by a high chief, 512.
name of, assumed by Kakuhihewa, 510.
placed in a canoe sent to Maui, 502.
receives harmless the spear thrusts of Kakaalaneo, 508.
rejecting ordinary food, ate bananas only, 500.
remains on Oahu with Kapuaokeonaona, 510–12.
repeats his visit to Hana and to Makolea from Kakaalaneo, 504.
replies to Makolea’s fears through her husband’s skill, 504.
returns unseen to Hawaii, 504.
says Makolea was taken by order of the king of Kauai, 510.
sets out for Kohala to return on third day, 506.
stands Kakaalaneo on his head, 504.
to enjoy Paliuli, 498.
with war club, cuts his opponent in two, 508.
with war club, meets Kakaalaneo holding two spears, 506.
with young wife on his back, retraces his steps, 508.
young wife, his foster-fathers and their wives in one canoe; the only one that
landed at Hana, 506.

Kiakia, bird-catching, 380.

Kiha, 25, 284, 302, 405.

Kihapaewa, or Kihapea, 336, 340.

Kihapiilani, advised to confer with Pao at Waikapu, 238.


advises wife of his departure, 242.
Aihakoko’s attendant killed by, 232.
and Piikea placed under Piilani, 236;
suggested as parents, 248.
arrives at Waipio; exchange greetings with his sister Piikea, and seeks Umi’s
aid, 244.
at Kalepolepo, sets out for Hawaii, 244.
beaten as he gathers potato tops; pays no attention, 238.
bids his discoverers “be quiet”, 238.
chief, unknown as such to the people, 236.
greets Pao and is instructed, 244.
neglected and ill-treated by Piilani; ran away secretly to Kalaniwai; marries
there, 236.
recognized as of high rank while getting potato tops, 238.
reveals himself, relates his ill-treatment and seeks for someone to avenge him,
236.
reveals his rank, 244.
sacred chief; a male through Piilani, 240.
searches for an avenger, 242;
seeks Pao, 244.
shall see bitterness, 240.
son of Piilani, chief of Maui, 242.
story of; to uplands of Kalaniwai, 242.
termed lazy by his wife’s parents, 236.
Umi turned Maui over to, 254;
went to the defence of, 232.
younger brother of Piikea, 236.

Kiholo and Kapalaoa, white sands of, 560.

Kii, 24, 404;


red rain of, 398.

Kiihele chides Kiinoho at his strange inaction, 502.


great runner, could circuit Hawaii in one day, 498.
questions Kiinoho who is to benefit by Paliuli’s delights, 498.
returns to Paliuli and reports his journey; narrates meeting with Makolea and
extols her beauty, 502.
sent to various districts for a suitable wife, 500–02.
takes Kepakailiula by the hand and leaves the house, 502.
told of Kiinoho’s dream; is indifferent and dreams same thing; traveler, 498.
unsuccessful till meeting Makolea in Kona, 500–02.

Kiinoho, a stay-at-home fortune-teller, 498.


bids Kiihele get Hina’s child, 500.
develops the egg in a feather cape into a beautiful child, 500.
dreams of Paliuli and tells Kiihele, 498.
sends Kiihele in search of wife for Kepakailiula, 500.

Kiinoho and Kiihele accompany Kepakailiula to Hana but not permitted to land,
506.
brothers of Hina, 498.
decide to find a wife for Kepakailiula, 500.
definition of, 498.
directed by dream, start for Paliuli, 498.
join in the fight, 508.
left Paliuli in charge of the gods, 502.
made joint kings of Oahu by Kepakailiula, reserving to himself and Kakuhihewa
rulers’ rights, 510.
mourn on leaving Paliuli, 502.
Puna chiefs of high rank, 498.
Kikakapu, butterfly-fish, 576.
put up in place of kapu stick, 576.
sacred fish, 240.

Kikenuiaewa, 24;
of Ewa, 342.

Kiki and party at Keolewa, 372.


hair dressing, 378.

Kila adjusts government of Kauai; declines the kingship, 152.


admired as a handsome young man, 134.
advised to delay departure, meets a priestess whose aid he invokes, 124.
again enters the temple, 144.
and brothers at Waipio, 132.
and Kamahualele seek in vain for place of Laamaikahiki’s hiding, 124.
and Laamaikahiki, arrival at Kauai, 128;
return to Tahiti with the bones of their father, 154.
and party set sail for Hawaii, 128.
anxious to find Laamaikahiki, 126.
arranges to take Moikeha’s bones to Tahiti, 154.
arrives at Luukia’s place and extends greetings, 124.
as Lena, questions Kaialea pointedly, to which false replies are given, 138.
asleep, is taken off the canoe and left at Waipio, 132.
assumes the reins of government on death of Moikeha, 128.
awakens and finds himself deserted, 132–34.
brings his mother and aunt into the temple, 148.
brothers questioned, orders them confined, 148.
bundled on the canoe platform, 164.
calls the people to witness the sacrifice, 148.
chants of Moikeha’s life of ease, 162.
chief ruler of Kauai, 152.
contends with Luukia, 172.
defers putting his brothers to death, 150.
did not think Kaialea would be killed, 148.
disregards mother’s desire that companions-in-death be offered up with their
sons, 150.
does not intend sacrifice, 144, 148.
does not wish the sacrifice of Kaialea, but his realization of the gravity of the evil
deed committed, 140.
drawn into a plot on pretext of brothers to obtain their father’s bones for removal
to Tahiti, 130.
elicited partial truth from Kaialea, 144.
falsely accused of violating kapus, flees to Pakaalana, 134.
follows the aged priestess; hides in the Mua house of the temple, 126.
former inhabitants wail on arrival of, 170.
[xxvi]gave up looking for Laamaikahiki; orders Kamahualele to prepare the canoe
for return, that Moikeha may send others, 124.
given the name of Lena, 134.
greets Laamaikahiki and is questioned, 128.
greets relatives at various points, 122.
handsome man, 164.
hearing of a canoe from Kauai, he sends for the men, 142.
Hooipoikamalanai and sister bewail the death of, 132.
informed of prayer’s interruption, 148.
insects, animals and the elements rejoice, at arrival of, in testimony of his high
chief rank, 168.
instructs a friend as to questioning Kaialea, 138.
instructs his men as to treatment of Mua; his wit avenges his father, 166.
instructs his men, in contest with two warriors, 166–68.
instructs the priests, on their course, 140.
Kaialea often questioned by, but found stubborn, 140.
Kaialea ordered confined again by, 144.
Kaialea search party questioned by, 142.
king and chief priest with, enter the kapu house, 148.
king of Kauai; jealousy of brothers thereat; reign of, not satisfactory, 130.
life of, in Waipio at first menial, 134.
lit the lamp and laid down on the couch, 170.
Luukia consenting, unfastens the cords, 172.
makes himself known, and relates his experiences, 150.
master of all the lands, victorious in his battle, 170.
meditates on his brothers’ actions, 134.
offers to die first; loved his brothers more than self, 152.
on the covered platform, 130.
orders food to be taken to Kauai for his mother and aunt, 140.
orders his officers to arrest the men, 142.
orders release of other men, 144.
orders reservation of food under penalty of death, 136.
originates the working system, 134–36.
otherwise known as Lena, 138.
prepares for the trip to Tahiti for Laamaikahiki, 120, 160.
prevailed on to be Kauai’s king, 152.
proclaims himself the offspring of Moikeha, 122–24, 128, 150, 162.
questions his mother; tells her Kaialea will surely die, 146.
questions Kamahualele, 124.
recognized by certain high chief signs, a priest directs the king of Waipio to take,
as a son, 134.
recognizes Kaialea’s canoe, and sees his brother; orders the canoe confiscated,
136.
recognizes Makalii, offers to meet him in contest, 168.
reported eaten by a shark, his hands only left, 132.
resembles Moikeha, 172.
restrained by his father from accompanying his brothers, 120.
retires to Lanikeha, 124.
returns to Waipio, 152;
to Kauai, 150.
returns to his father’s house; the guards come to life, 170.
reveals himself and the object of his journey, 126.
sails for main island, 164.
saw his mother and relatives; orders houses made ready, 146.
seen by Luukia asleep, is mistaken for Moikeha; embraces him, is startled, 172.
sends men to take Kaialea to the king’s strong house, 138.
sets sail for Oahu on voyage to Tahiti, 122.
son of Moikeha, 118;
and Hooipoikamalanai, 160.
spared from death through intervention, 134.
standing by the anuu, faces his brothers, 148.
suggests a god be provided his brothers, 120.
suggests to his mother and aunt that Umalehu and the rest be saved; is
opposed by them, 150.
taken by the brothers, they sail for Oahu, 130.
tells his men to return to Kauai if he is slain in contest with Makalii, 168.
tells history of his brother’s treatment, 134.
the crowd shout in praise at sight of, 168.
the shells advise there are no more chiefs, 170.
told of the food delivery to his people, whereas it was all squandered at Molokai,
140.
told of Kaialea’s weeping, he questions him, 144.
tries to conceal his emotions, 140.
upon death of Moikeha the land descended to, 128.
visits and exchanges greetings with Kanepohihi, 162.
wins in contest with his brothers, 120, 160.

Kila’s brothers dissuade the mothers from joining, 130.


fear their scheme will fall through, 130.
hear there is food at Waipio, 136.
kidnap a young man from Waipio and slay him, 132.
plan concealment of their jealousy and hatred; professing obedience, plot
against him, 130.
prepare the double canoe, planning to include Kila, 130.
propose to bring the bones of their father for removal to Tahiti, 130.
report Kila as eaten by a shark and bones of father lost, 132.
swearing to take good care of Kila, the mother’s fears are allayed, 130.
take Kila and sail for Oahu, 130.

Kila-pa-Wahineikamalanai, 122–24, 128, 150.

Kilauea, pit at, dug by Pele and Hiiaka, 106.


to Kalihi, 358.

Ki-leaf (or Ti-leaf) fishing coat, 224;


knotted, 366.

Kilohi begs Wahanui to return to Hawaii, 518.


not the priest Wahanui had thought him to be, 518.
prophet of Wahanui, joins him on voyage to Tahiti, 516.
refuses Kaneapua to board their canoe, 516.
terms Kanehunamoku the man-eating dog of Hina, 518.

Kilou, cliff of Lehua at, 306.

Kinau, a sand-eel, 358.

King of Hawaii, Kapawa, 22.


of Kauai, Ku, 372.
of Kauai meets Kualii and gives over his island to him, 400.
of Koolau (Kualii), 402.
of Koolauloa ceded the districts to Kualii, 400.
of Kauai, Moikeha became, 118;
Kiha made, 130.

King’s loin cloth and kapa, 278.

King’s riddles, Kepakailiula given the answer to, 512–14;


invited to join in solving, 514.
Kepakailiula solving the; Kukaea throws the king into the oven, 514.
Kukaea summons the people to answer the, 512.
oven-baking the penalty in contest of, 514.

Kini (40,000), 364;


from word Kinikini, 400.

Kinilauaemano, 370, 404.

Kino, or miraculous powers, 72.

Kio, 24, 404.

Kipahulu, Koi and companion sail from, 234.

Kipapai, 288, 304.

Kipapalaulu, asked by daughter, sends Aiai a ten-fathom canoe, 558.


asked for a pearl fish-hook, 556.
[xxvii]king of Honolulu, at success of Kuula in aku fishing steals his pearl hook
Kahuoi, 556.
living at Kapuukolo, 546.
on further request by daughter, gives up the stolen hook Kahuoi, 556.

Kipu, mischief-maker of Palaau, 396.

Kipunuiaiakamau and companion on Moikeha voyage from Tahiti, 116.


navigators and sailing masters with Kila, 122.

Kiss on the nose, 350.

Kissing of olden time, “honi ka ihu”, 308.

Kiu ahiu, wild spy, 396.

Kiwaawaa, a coarse kapa, 584.


Kiwaha gives Aukele a way of escape by rainbow, 66.
mate of Halulu, 66.
one of three bird-guards of Namakaokahai, 42.

Kiwalao, overthrow of, 4.

Koa, 25, 405.

Koa tree, sounding-leafed, 358.


trunkless, 350–52;
without roots, 356.

Koae (bird), 70, 234, 340;


that soars high, 394.
bos’n bird (Phaethon lepturus), 340, 394.

Koaie (tree) of Kauai, 386.

Koakea, 186;
heights of, adjoining Waipio, 208.
Umi meets Piimaiwaa at, 182.

Koauli, chant, 342.

Koeleele powerful man of bad temper, 528.


runs away from Kaulu, 528.
younger brother of Makalii hurls the rock Ikuwa at Kaulu, 528.

Kohala, aeloa the favorite wind of, 566.


ahupuaa of Hihiu nui in, 354;
Kapaihiahilina lands at, 356.
and Hamakua rebels met at Nakikiaianihau, 324.
beautiful country, 314.
bones of chief of, 314;
boundary between Kona, 362.
depopulated, 336–38.
did not see people of, 338.
dividing line of, 360.
east of, without growing food, 570.
forsake the proud land of, 568.
given to Koi, 206.
got their fish from Mumu and brothers, 562.
heiau of Muleiula in, 324;
in Kapaau of, 328.
is in darkness, 28.
Kaiopihi, the slain general, 330.
Kapua and Kukuipahu in, 380.
Kauhiakama arrived at, 334;
reports on his tour through, 336.
lies level, face down, 314.
Lono and forces reach; victory followed into, 328.
lowland country noted as a proud land, 540.
making and worshipping of idols originated in, 540.
men all at, awaiting slaughter by Lono, 320.
men had also been assembled and guarded, 344.
of Wakiu, 306.
Palahalaha, chief of, 314.
people of, attached to, 540.
Puuonale, king of, 538.
rebels stationed from Anaehoomalu to distant, 322.
small district, to be fought last, 324.
stretches forth, 374.

Kohana, naked, 378.

Kohenemonemo, wife of Hauna and Loli, 256.

Kohia, from Ko, 372.

Kohikohiokalani, 24.

Koholalele, Liloa journeys to, 178;


Umi and companions journey to, 186.
shallow sea of, 398.

Koi accompanies Umi to Laupahoehoe beach, 210.


and co-counselors ordered to prepare canoes, 246.
and companions with the king reside at Waipio, 214;
become courtiers, 220.
and companions in spear practice with Umi, 210.
and companions without bundles of stones, 200–02.
and officers ordered to war on stronghold of Kauiki, 248.
at call of Piimaiwaa followed him up, 254.
brother-in-law and, select Umi’s burial place, 234.
charged with secretion of Umi’s bones at his death, 232.
directed to kill Paiea; returns with the body for sacrifice, 214.
dispossessed of his lands, 232.
enters home of his sister, 234.
finds the guards asleep, takes the body of Umi, 234.
hearing of Umi’s fatal illness, sets out for Hawaii, 232.
hears of Umi’s death, 234.
kills a man as a substitute corpse for Umi, 234.
met at Kukuihaele and taken along by Umi, 186.
Omaokamau and Piimaiwaa aku fishing; farm daily, 186;
taught the arts of warfare, 190.
recognized, enjoins quiet and secrecy, 234.
report of, makes Umi sad at heart, 250.
said to have taken Umi’s bones to Maui, 234.
sees Umi’s hurt, vows to slay Paiea, 210.
sent to scale Kauiki; after two attempts, in fear of the giant guard he returns,
250.
Waimanu to Pololu the lands of, 232.
wanders away to Kauai, 232.
with Umi and Omaokamau, confined for sacrifice, 220.
with valuables, sails secretly for Maui, 234.

Koihalawai, 25, 405.

Kokio, Ku not like the, 392.


medicinal, 390.

Koko, a net, 530.


Kualii held up in his, 388;
network, 364, 400.

Kokoiki, Kohala, Maile sisters met Hikapoloa at, 562.

Kolea and mate fly up and inquire of Makalii of the loud-voiced god Kaeha, 524.
reports Makalii’s message, 524.
said to Ulili, “Let us fly high above Kana and call to him”, 444.
told of Kaulu hiding in the palm leaf, 524.
Kolea and Ulili are told by Hina wherein Niheu’s strength lies, 446.
fall down on the hill of Haupu, 444.
met Kapepeekauila, barely escaped death; sent to tell Keauleinakahi, his
warrior, 444.
not a formidable pair, 444.
seeing Hina being taken, flew and held Niheu by the hair, 446.
swift messengers of Kapepeekauila, sent to look for Kana and Niheu, 444.

Kolohia, 374.

Kolowalu (Royal) Statute, a beneficent law of Kualii, 432.

Kona (Oahu), 300, 384;


another wing of army from, 410.

Kona and Kau, kukui first introduced into, 570.

Kona and Koolau (Molokai) continue the battle against Kekaha, 418.
bones of chief of, 320;
rebels from, arrive, 330.
district, defeated king of, 394.
district, the largest, 338.
dividing line of, 360–62.
dwell in; house stands in, 286, 304.
Ehunuikaimalino king of, 228.
first meets the eye, 374.
given to Ehu, 206;
Kapalilua, 336.
Heapae chief of, 320;
Lono at temples in, 330.
Kapaihiahilina sails for, 356;
returns to, 362.
Kauhi through, sees not its people, 338.
Kauhiakama reports on, 336.
[xxviii]known from below, 378.
men from, 344.
Moihala chief of, son of Heapae, 320.
stands forth to sight, 28;
plainly seen, 374.
term for the lower regions, 378.
the sun warmed the selfish chiefs of, 394.
Umi desired to live in, 228–30.
whose stone floor burns, 394.

Konane board, Lono strikes his wife with, 272.


chant, 56, 272;
quoted in contests, 272.
engaged in playing, Lono strikes Pupuakea on the head with the board, 334.
game of, 56, 270–72;
resembling checkers, 270.
Hauna plays, against two women, wagering his canoe load of feathers and wins,
312.
Kakuhihewa challenges Lono to a game of, 300.
Kama and Lono played, 334.

Koniewalu, 370, 404.

Konohiki, 24, 404.

Kookookumaikalani, 25, 404.

Koolau, 284–86, 300, 304, 392;


army from, 410.
bracing up the house of, 392.
chief of, 366;
flower of, 314.
Lonokaeho, king of, 530.
tea plant (Campylotheca), 386.
trembles, 396.

Koolau and Kona (Molokai) chiefs battle against those of Kekaha, 418.
chiefs of, gave up to Kualii all Molokai, 420.
chiefs of, hear war is to be carried into Kalaupapa, 418.
defeated, lands on the, side come into Paepae’s possession, 418.
war canoes from all the side of, go to battle, 418.

Koolaukahili, 25, 405.

Koolauloa, 300, 364, 388.


armies of chiefs of Waianae and, routed, 414
and Koolaupoko ceded to Kualii, 400.
Koolaupoko, Kailua in, Kualii’s favorite residence, 432.
Kualii and boy returned to Kailua in, 430.
Kualii died at Kailua, in, 432.

Kou, Honolulu harbor, 452–54.


Kahaookamoku’s party lauded at, 478.
scattering blossoms of the, 392.

Koukou, drank the awa of, 378.


mother of Pikoiakaalala, a son, and Iole and Opeapea born before him, 450.

Kowali or Koali, 530.

Kowili, 372.

Ku (deity), 26, 30, 394.

Ku enamored of Kaunoa while bathing, 548.


father of Kalanimanuia, 548.
king of Lihue, takes Kaunoa to wife, 548.
leaves Kaunoa with expectant child, 548.
leaves name and tokens for the child, 548.
not recognizing the lad, orders him killed, 548.
on hearing strict kapu violated, orders his female attendant killed, 542.
on priests’ questions, owns the relationship with Kaunoa, 550.
realizing having killed his own son, seeks to regain him, 550.
searched for by Kalanimanuia, 548.
thou unnatural father, 548.

Ku (Kualii), 30, 372, 376, 380, 386, 390–96, 414–16, 420, 428.
arrayed in his feather cloak, 384, 416.
encompassed by, is the island, 400.
haole from Tahiti, a god, 394.
has left but few priests, 386, 416.
holds up the rain, 378;
led to earth, 380.
indeed, whose is Tahiti for, 374.
is brought forth in the forest, 384.
is indeed king, 384–86, 416.
puts on his loin-cloth for war, 382.
returning to Oahu; sailing to Kauai, 374.
the lehua eater, 286, 304.
uncomparable, 390–94.
urged to be merciful and spare his wrath, 388.

Ku and Hina, male attendant reports the conduct of their two charges to, 542.
parents of Kepakailiula, 498, 540.
son and daughter of, brought up under strict kapu, 540.

Kua, maile vines creep down to, 400.

Kuaihelani, 32, 46–48, 68.


Aukelenuiaiku the hero of, 78;
his departure for, 108.
Bambu stalk shoots up till it reached, 598.
boy from, 56, 80.
brother or sister to be banished to, 540.
brothers of Aukele depart for, but meet disaster and perish, 106.
champions, visit Kauai, Oahu, and Maui; give exhibition of games, 34.
deserted and overgrown with weeds, 108.
fine rain of, 94.
Hina originally belonged to, 546.
home of Makiioeoe, where, as king, he had one child, Kahikiula, 596.
Kanemoe accompanies Aukele to, 108.
Kapuaokaoheloai and messengers embark for, 542.
king of, desires to know his daughter’s opinion, 544.
king of, questioned the priests relative to rank of Kapuaokaoheloai, 544.
land in Tahiti, 540.
land of, origin of Aukelenuiaiku, 32.
Laukiamanuikahiki told her father has gone back to, 596;
said “Here I am returning to,” 608.
Makiioeoe prepares and returns to, 596.
messengers of, cautioned Kapuaokaoheloai regarding the king’s daughter, 542–
44.
messengers of king of, in search of a wife, 542.
popular mythical land, 32.
the chief reminded that banishment to, would be the penalty for violating the
kapu, 542.

Kuaikalolo and Aiakoake, elders of Kepakailiula, 498.


Kuaiku, 32, 36;
has his arms broken, 36.

Kuaimakani, 180.

Kuaiwa, 25, 405.

Kuaiwilu, a voyager with Kila to Tahiti, 122.

Kuala, current of, 240.

Kualii, 364–66, 370.


accompanies his soldiers in battle usually, 426.
advised by priest of Pumaia’s spirit as the thief, 476.
and Maheleana take war club lessons, 412.
and warriors sail to Kauai for war clubs; armies different, closing in on, 412.
arrival of, reported to Haloalena and Kamalalawalu, 424.
arrives at Kahaluu, 428.
arriving at Laupahoehoe, Haalilo prepares for war, 414.
asks Kauhi’s authority for his mischievous acts, 426.
assumes a royal right to dedicate Kawaluna temple, 408.
astrologers fail to find auguries for defeat of, 366.
at advice of priest, builds houses and cares for bones of Pumaia, 476.
awakens his companions to meet the advancing armies; refuses to flee, 408–10.
battle of Kalena, names it the, 414.
battles and battle grounds of, 406;
fights three more battles, conquering Oahu, 414.
began fighting in childhood; story of, 364.
bestows great riches and favors on Kapaahulani; orders share sent the brother
at Puuloa, 402.
body of Pumaia thrown into pit of temple of, 472.
[xxix]bones of, to be secreted at death by his trusted kahu, are powdered and
hidden in one hundred living tombs, 434.
calls and questions the boy, deeming him very brave, 430.
celebrated for strength and bravery, 364.
chant composed to name of, 364–66;
supplementary chant for, 394.
charged with having overstepped himself, 408.
chief officer of, remarked, 378.
compared to a god and an early king, 388.
declares the battle prepared by Haloalena off, 426.

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