SPECIALISSUE

The maximum age of Hawaiian terrestriallineages: geological constraints fromK�oko Seamount

David A. Clague1*, Juan C. Braga2, Davide Bassi3, Paul D. Fullagar4,

Willem Renema5 and Jody M. Webster1,6

1Monterey Bay Aquarium Research Institute,

7700 Sandholdt Road, Moss Landing, CA

95039-9644, USA, 2Departamento de

Estratigrafıa y Paleontologıa, Universidad de

Granada, Campus Fuentenueva, E-18002

Granada, Spain, 3Dipartimento di Scienze

della Terra, Universita di Ferrara, Via Saragat

1, I-44100 Ferrara, Italy, 4Department of

Geological Sciences, University of North

Carolina, Chapel Hill, NC 27599-3315, USA,5Nationaal Natuurhistorisch Museum, Leiden,

The Netherlands, 6School of Geosciences,

The University of Sydney, Sydney, NSW 2006,

Australia

*Correspondence: David A. Clague, Monterey

Bay Aquarium Research Institute, 7700

Sandholdt Road, Moss Landing, CA 95039-9644,

USA.

E-mail: clague@mbari.org

ABSTRACT

Aim To determine if K�oko Seamount submerged below sea level before Kure

Island and Pearl and Hermes Reef formed, resulting in a period in which there

were no extant islands. A period with no islands would eliminate prior terrestrial

and shallow marine biotas that could migrate from island to island and require a

restart of colonization from distant shores to populate the younger islands of the

Hawaiian volcanic chain.

Location Emperor Seamount Chain, north-central Pacific Ocean.

Methods We estimate subsidence rates for K�oko Seamount using ages

determined from fossil large foraminifera and Sr-isotopes, and maximum

depths using palaeodepth estimates based on coralline algae. These data are

combined with palaeolatitude changes as the Pacific Plate moved northwards, sea

level variations, and sea surface temperature variations at the seamount through

time to reconstruct the time and causes of submergence.

Results Rounded carbonate clasts include three facies: zooxanthelate corals,

bioclastic packstones to rudstones, and rhodolith floatstones. Two rudstones

contain relatively deep-water, coralline algal rhodoliths and large foraminifera

indicative of Aquitanian (20.4–20 Ma) and Burdigalian (20–16 Ma) stages of the

Early Miocene, consistent with Sr-isotope ages of algae and one sample of large

foraminifera. Corals grew on K�oko Seamount from c. 50 to 27.1 ± 0.4 Ma, the

youngest Sr-isotope age of a coral sample. These shallow, warm-water coral reefs

came under increasing stress as the volcano subsided at 0.012 ± 0.003 mm yr)1,

and migrated northwards, and as global climate cooled. The summit submerged

and shallow coral reef growth ceased before 29 Ma, probably around 33 Ma. The

volcano continued its slow subsidence, and deep-water carbonates accumulated

until they too were unable to keep pace, dying out at c. 16 Ma.

Main conclusions The final submergence of the summit of K�oko Seamount by

about 33 Ma confirms that biota on older Hawaiian–Emperor Islands could not

have migrated from island to island along the entire chain to eventually colonize

the present Hawaiian Islands. There was a period between at least 33 and 29 Ma

in which no islands existed, and distant colonization had to repopulate the

younger portion of the Hawaiian chain, which began to emerge between about 29

and 23 Ma.

Keywords

Bathymetry, carbonates, Emperor Seamounts, Hawaii, island biogeography,

submergence, subsidence.

Journal of Biogeography (J. Biogeogr.) (2010) 37, 1022–1033

1022 www.blackwellpublishing.com/jbi ª 2009 Blackwell Publishing Ltddoi:10.1111/j.1365-2699.2009.02235.x

INTRODUCTION

Age estimates of most recent common ancestors (MRCAs) of

lineages of Hawaiian organisms (summarized in Price &

Clague, 2002) show that 19 of 22 lineages originated following

the formation of Kauai. However, three important lineages are

inferred to have originated earlier: fruitflies at 26 Ma (Russo

et al., 1995), lobeliods at 13 Ma (Givnish et al., 2009) and

damselflies at 9.6 Ma (Jordan et al., 2003). These constitute

evidence of an evolutionary ‘conveyor belt’ (Carson, 1983)

prior to the formation of Kauai. However, Carson & Clague

(1995) and Clague (1996) argued, based on subsidence models,

that all previously formed islands had submerged by the time

that Kure Island formed c. 30 Ma, possibly providing a

geological explanation for why no older lineages have been

found.

The evolution of the Hawaiian–Emperor volcanic chain

(Clague & Dalrymple, 1987) is well known, as are the ages of

many volcanoes in the chain (summaries in Duncan & Keller,

2004; Sharp & Clague, 2006). In contrast, relatively little is

known of the subsidence histories or longevities of former or

extant islands. Such data are important for understanding the

sequential colonization of each new island because most

terrestrial and shallow-marine biotas originated from pre-

existing, nearby islands in the chain (Carlquist, 1995).

Clague (1996) used bathymetric data and a simple subsi-

dence model to estimate the areal extent and heights of the

islands along the chain, and estimated the life span of each

island. His model suggested that a significant barrier to

terrestrial species migration existed so that islands in the chain

younger than 30 Ma were colonized from distant shores,

because no two islands existed simultaneously from 33 to

30 Ma. K�oko Seamount, in the southern Emperor Seamounts

(Fig. 1), figures prominently in this biogeographic model of

the chain because it was large and therefore survived as an

island for longer than any of the other volcanoes older than

17 Ma. The model for K�oko Seamount indicated that it

submerged about 36.5 Ma, 14 Myr after the volcano formed at

50.4 Ma (Sharp & Clague, 2006) and at least 7 Myr before the

next large island (Kure Atoll) grew (Clague, 1996).

We present new age and palaeoenvironmental data from

some shallow-water carbonate samples from the summit of

K�oko Seamount. The facies and ages allow us to determine the

subsidence history and submergence age of the island and to

determine if the geological history of this key seamount is

consistent with the MRCA analyses of Hawaiian lineages.

GEOLOGICAL SETTING AND PREVIOUS WORK

K�oko Seamount is in the southern Emperor Seamounts, near

the bend of the Hawaiian–Emperor chain, at c. 35�20¢ N,

171�30¢ E in the northern Pacific basin (Fig. 1a). This chain

consists of at least 129 volcanoes, comprising a hotspot track

formed as the Pacific plate has moved over a mantle magma

source over the past c. 80 Myr (Wilson, 1963; Clague &

Dalrymple, 1987; Sharp & Clague, 2006). K�oko Seamount rises

from the abyssal plain at c. 5500 m to as shallow as 253 m

below sea level (Fig. 1c). It is a broadly elliptical guyot, or flat-

topped seamount, aligned in a NNW–SSE direction, with a

slightly domed summit plateau at c. 350 m depth (Davies

et al., 1972). The bathymetry suggests that the Seamount is

composed of at least three large, coalesced volcanoes

surmounted by a coral reef or bank (Fig. 1c).

The summit has been imaged using seismic reflection

profiles, which reveal a carbonate cap (Davies et al., 1972).

Using additional seismic reflection profiles, Greene et al.

(1980) interpreted the summit as having an extensive carbon-

ate cap with patch reefs and an ancient inner lagoon covering

about 650 km2. The largest reef structure was mapped in the

south-east part of the summit, upslope from our sample site,

and the lagoon was located to the north-northwest, near the

centre of the summit. Greene et al. (1980) inferred that the

base of the carbonate deposits was at 785 m depth, indicating a

thickness of about 530 m.

Deep Sea Drilling Program (DSDP) sites 308 and 309

(Shipboard Scientific Party, 1975a,b) and Ocean Drilling

Program (ODP) site 1206 (Shipboard Scientific Party, 2002)

drilled K�oko Seamount (locations shown on Fig. 1c). The

cores from sites 308 (depth 1331 m) and 1206 (depth 1557 m)

recovered shallow-water large foraminifera, coralline algae,

green algae (Halimeda), bryzoans and coral fragments that are

from NP14 or NP15 zones in the Early to Middle Eocene

(43.4–52.4 Ma; Matter & Gardner, 1975). At site 309 (depth

1454 m) a tiny fragment of cuttings included large foraminif-

era of Late Oligocene to Early Miocene (c. 28.4–16 Ma) age,

with coral fragments and coralline algae. The youngest known

carbonates, prior to this study, are at least 8 Myr younger than

the model subsidence age of 36.5 Ma from Clague (1996).

Isotopic (40Ar/39Ar) ages of shield and post-shield volcanic

cobbles dredged from the southern part of K�oko Seamount

(D43, shown in Fig. 1c) yield a best age of 50.4 ± 0.1 Ma in

the Early Eocene (Sharp & Clague, 2006). Some zooxanthelate

corals recovered with the volcanic cobbles were examined by

W. Durham at the University of California at Berkeley, who

found them to have a maximum age of Lower Tertiary, no

older than Palaeocene (quoted in Davies et al., 1972). Ar–Ar

isotopic ages on shield-stage lava flows recovered at ODP

site 1206 (Duncan & Keller, 2004) are slightly younger

(49.1 ± 0.2 Ma) than the Sharp & Clague (2006) age for

southern K�oko Seamount. The fossil samples are coeval with

or younger than the isotopic ages of the dredged volcanics and

suggest that the carbonate cap accumulated from near the time

of volcano formation at 50.4 Ma until sometime between 28.4

and 16 Ma.

Palaeomagnetic data from ODP site 1206 suggest that K�oko

Seamount formed at about 21.5� N during the Palaeocene

to Middle Eocene, a period of southward migration of the

Hawaiian hotspot (Tarduno et al., 2003). By c. 45 Ma, the

hotspot was no longer migrating southwards and was situated

at about 19� N, its present latitude.

Bioclastic sediments on Suiko (DSDP site 433 during Leg

55), Nintoku (site 432), and Ojin (site 430) seamounts to the

Geological constraints of Hawaiian lineages

Journal of Biogeography 37, 1022–1033 1023ª 2009 Blackwell Publishing Ltd

(a)

(b)

(c)

(d)

(e)

(f)

(g)

Figure 1 Bathymetric maps of K�oko Seamount. (a) Regional map of the north-central Pacific Ocean showing the Hawaiian–Emperor

volcanic chain. The box labelled b is the extent of a closer view of the western Hawaiian Ridge shown to the right in panel (b). In panel (b),

the box labelled c is the extent of the map of K�oko Seamount shown in panel (c). These maps are based primarily on satellite altimetry

data: Geoware GMT Companion CD-R, Vol. 1, version 1.9, June 2006. The 30-s blend of observed and predicted topographic data are

gridded at about 925 m and shown with 500-m contours and a colour scale from deep blue at 6000 m to pale orange at sea level. The

locations of D43 from the Aries VII cruise, and Deep Sea Drilling and Ocean Drilling Program sites 308, 309 and 1206 are shown.

The extents of (d), (e), (f) and (g) are shown in (c) and are based on a single multibeam swathe from JAMSTEC cruise KR03-10. The data are

gridded at 50 m using MB-System software (Caress & Chayes, 2004) and are shown as slope-illuminated maps. The scale bar beneath

each letter is 1 km long. Panels (d) and (e) show the steep eastern and western break-in-slope at c. 1555 m depth and a series of ridges

and terraces above the break-in-slope that resemble coral reef structures seen around Hawaii using a colour scale from deep blue at 2500 m to

pale orange at 1250 m. Panels (f) and (g) show terrace scarps with vertical offsets up to 150 m tall using a colour scale from pale orange

for 300 m to deep blue at 1050 m.

D. A. Clague et al.

1024 Journal of Biogeography 37, 1022–1033ª 2009 Blackwell Publishing Ltd

north of K�oko Seamount consist primarily of coralline algae

and bryozoans with ostracods, foraminifers and assorted

mollusc shell fragments typical of shallow-water, high-energy

environments (summary in Jackson et al., 1980). In the

modern north Pacific, coral–algal facies dominate below 20�latitude, and algal–bryozoan facies dominate at latitudes above

30�, with the boundary at c. 25� (Schlanger & Konishi, 1975).

The algal–bryozoan facies bank deposits on these seamounts

were used to argue for the northward migration of the Pacific

Plate during the formation of the Emperor chain (Jackson

et al., 1980). The only other carbonate bioclastic sediment

recovered from the southern Emperor Seamounts was a single

clast containing large shallow-water foraminifers of Late

Eocene age dredged from Kammu Seamount at the southern

end of the chain (N. Sachs, Smithsonian Institution, pers.

comm., quoted in Clague & Jarrard, 1973).

MATERIALS AND METHODS

Samples were collected by dredging at the south-eastern edge

of the summit plateau in August 1971, on Leg 7 of the Scripps

Institution of Oceanography’s ‘Aries’ Expedition on the

Research Vessel Thomas Washington. Dredge 43 was on

bottom over a depth interval from 827 to 628 m (Fig. 1c).

The haul recovered a wide range of volcanic cobbles, including

phonolite, trachyte, benmoreitie, mugearite, hawaiite, alkalic

basalt and tholeiitic basalt (Clague, 1987), as well as limestone

pebbles to cobbles with corroded and bioeroded surfaces

(Davies et al., 1972). Thirty-two of these carbonate samples

were cut to identify lithofacies and macrofossil components,

and thin sections were made to identify microfossils, especially

coralline red algae and benthic foraminifers. The fossil

components are listed in Table 1.

Twelve samples for 87Sr/86Sr analyses were ultrasonically

cleaned in deionized water and dried. The most pristine (lack

of obvious macroscale secondary precipitates, mud and/or

discoloration) part of each sample was selected under a

petrographic microscope, but several of the coral samples

have clear evidence of recrystallization. A 1-mm drill was

used to precisely subsample corals, coralline algae crusts and

foraminifera-rich sediments. 87Sr/86Sr measurements were

undertaken at the University of North Carolina Department

of Geological Sciences using a VG Sector 54 mass spectrom-

eter. The ratios are reported with 2r errors relative to the87Sr/86Sr ratio of 0.710250 for the carbonate standard SRM

987 (Table 2).

RESULTS

Sedimentary facies and age data

Four samples (A5, A6, A19 and A30), from 5 to 10 cm across,

are made up entirely of colony skeletons of zooxanthelate

corals: Porites and unidentifiable faviids (Fig. 2a). Samples A12

and A19 are locally encrusted by coralline algae. Only

Lithophyllum gr. pustulatum can be identified among the

coralline algae. Sample A18 is mainly a coral clast set in a

rudstone with abundant large benthic Amphistegina foramini-

fers in the matrix. Coral fragments also occur in A29, another

rudstone (Table 1). Four of five analysed corals have ages

based on Sr-isotopes ranging from 27.1 to 29.9 Ma, or from

Rupelian to Chattian stages in the Oligocene (Table 2). Sample

A30 yielded a Pleistocene age, which is considered erroneous.

Coralline algae surrounding a coral clast in A19 has an

age based on Sr-isotopes of 28.15 Ma, consistent with the

coral ages.

The bioclastic packstones and rudstones facies consist of

fragments of bryozoans, coralline algae (Fig. 2b), molluscs and

larger benthic foraminifera with a muddy matrix. Small coral

fragments, serpulids, echinoids and small benthic and plank-

tonic foraminifera can be additional, minor components

(Table 1). Many skeletal particles have dissolved, leaving voids

in the matrix, and fragments in several samples have recrys-

tallized and cannot be identified. Dissolution has preferentially

affected aragonitic skeletons. Amphistegina (Fig. 2b) is the

most abundant larger benthic foraminifer. Amphistegina can

account for more than 80% of skeletal components in a few

samples. Coralline algae are represented by encrusting and

loosely branched plants of melobesioids [Lithothamnion

(Fig. 2d) and rare Mesophyllum]. Eight of 12 packstones do

not contain coralline algae. The coarse fragments in most

rudstones are algae; those with concentric arrangement are

similar to the rhodolith floatstone facies.

The rhodolith floatstone facies comprises coralline red

algal nodules (rhodoliths) embedded in a packstone matrix

(Fig. 2c). Components in the matrix are mainly small benthic

foraminifers and bryozoan fragments. The benthic foraminif-

era Lepidocyclina (Euepidina), Neorotalia and Spiroclypeus

(Fig. 2e) occur in one sample together with rare Amphistegina

and Operculina. Rhodoliths are ellipsoidal to spherical and

up to 7 cm in maximum dimension. They are made up of

encrusting, warty and lumpy plants of Lithothamnion and

Sporolithon, and rare Lithophyllum gr. pustulatum, intergrown

with scarce laminar bryozoans (Fig. 2d).

Five samples of algal crusts from rhodoliths yielded ages

based on Sr-isotopes that range from 28.8 to 21.2 Ma, namely

from the Rupelian stage of the Oligocene to the Aquitanian

stage of the Early Miocene. A single sample of Amphistegina-

rich sediment yielded an age based on Sr-isotopes of

16.2 ± 0.2 Ma, namely the late Burdigalian stage of the Early

Miocene.

Bathymetric data

The main break-in-slope on K�oko Seamount is at c. 1555 m

(identified from a single JAMSTEC (Japan Agency for Marine-

Earth Science and Technology) SeaBeam swathe profile in

Fig. 1d–g), which implies that this much subsidence has

occurred since volcanic activity stopped repaving the shoreline

(Moore & Clague, 1992) at c. 50 Ma. This depth to the break-

in-slope is greater than that around most of the main Hawaiian

Islands (generally between 1000 and 1100 m).

Geological constraints of Hawaiian lineages

Journal of Biogeography 37, 1022–1033 1025ª 2009 Blackwell Publishing Ltd

Tab

le1

Fac

ies

and

carb

on

ate

com

po

nen

tso

fK

� oko

Seam

ou

nt

sam

ple

s.

Sam

ple

*F

acie

sL

arge

fora

min

ifer

aB

iost

rati

grap

hic

age

Co

rall

ine

alga

eP

alae

od

epth

Oth

erco

mp

on

ents

A1

Pac

ksto

ne

Lit

hot

ham

nio

nsp

ecie

s>

20m

Bry

zoan

s,p

lan

ktic

fora

min

ifer

s

A2

Ru

dst

on

eT

wo

dif

fere

nt

Lit

hoth

amn

ion

spec

ies

>20

mC

ora

l,m

ollu

scfr

agm

ents

,b

ryo

zoan

s,p

lan

ktic

fora

min

ifer

s

A4

Ru

dst

on

eT

hre

ed

iffe

ren

tL

itho

tham

nio

nsp

ecie

s>

20m

Bry

zoan

s,se

rpu

lid

s?,

ben

thic

fora

min

ifer

s

A7

Rh

od

oli

thL

ith

oth

amn

ion

spec

ies

>20

mB

ryzo

ans,

oys

ter

frag

men

ts

A8

Rh

od

oli

thA

mph

iste

gin

a,tw

o

dif

fere

nt

Ope

rcu

lin

a,

Ope

rcu

lin

ella

Bu

rdig

alia

n

A9

Pac

ksto

ne

Am

phis

tegi

na

Mel

ob

esio

ids

ind

et.

>20

mB

ryzo

ans,

ech

ino

ids,

pla

nkt

ican

dsm

all

ben

thic

fora

min

ifer

s

A10

Ru

dst

on

eA

mph

iste

gin

aL

ith

oth

amn

ion

spec

ies

and

mel

ob

esio

ids

ind

et.

>20

mB

ryzo

ans,

pla

nkt

ican

dsm

all

ben

thic

fora

min

ifer

s

A11

Pac

ksto

ne

Am

phis

tegi

na

Bry

zoan

s

A12

Co

ral

Lit

hop

hyl

lum

gr.

pust

ula

tum

Alg

aen

ot

ind

icat

ive

Bry

zoan

s,ec

hin

oid

s,m

ollu

scs

A13

Ru

dst

on

eM

elo

bes

ioid

sin

det

.>

20m

Bry

zoan

s,sm

all

ben

thic

fora

min

ifer

s

A14

Pac

ksto

ne

Am

phis

tegi

na

Bry

zoan

s,m

ollu

scs,

smal

lb

enth

icfo

ram

inif

ers

A15

Pac

ksto

ne

Am

phis

tegi

na

Spor

olit

hon

gr.

mol

le>

50–

60m

Bry

zoan

s

A16

Pac

ksto

ne

Mo

llu

scfr

agm

ents

A17

Pac

ksto

ne

A18

Ru

dst

on

eA

mph

iste

gin

aL

arge

cora

lfr

agm

ent

A20

Rh

od

oli

thT

wo

dif

fere

nt

Spor

olit

hon

spec

ies,

Lit

hoth

amn

ion

spec

ies,

Lit

hoph

yllu

mgr

.pu

stu

latu

m

>50

–60

mSm

all

ben

thic

fora

min

ifer

s

A21

Ru

dst

on

eA

mph

iste

gin

aM

elo

bes

ioid

sin

det

.>

20m

Bry

zoan

s,ec

hin

oid

s,m

ollu

scs,

smal

lb

enth

icfo

ram

inif

ers

A22

Rh

od

oli

thL

ith

oth

amn

ion

spec

ies,

Spor

olit

hon

spec

ies,

mel

ob

esio

ids

ind

et.

>50

–60

m

A23

Rh

od

oli

thA

mph

iste

gin

a,Sp

iroc

lype

us,

Lit

hot

ham

nio

nsp

ecie

san

dm

elo

bes

ioid

sin

det

.>

20m

Bry

zoan

s,ec

hin

oid

s,m

ollu

scs,

smal

lb

enth

icfo

ram

inif

ers

Lep

idoc

ycli

na

(Eu

lepi

din

a),

Neo

rota

lid

s

Aq

uit

ania

n

A24

Pac

ksto

ne

Am

phis

tegi

na

Mo

llu

scfr

agm

ents

A25

Pac

ksto

ne

Am

phis

tegi

na

A26

Rh

od

oli

thA

mph

iste

gin

aM

elo

bes

ioid

sin

det

.>

20m

Bry

zoan

s,m

ollu

scs,

smal

lb

enth

icfo

ram

inif

ers

A27

Pac

ksto

ne

Am

phis

tegi

na

Mel

ob

esio

ids

ind

et.

>20

mB

ryzo

ans

A28

Pac

ksto

ne

Bry

zoan

s

A29

Ru

dst

on

eA

mph

iste

gin

aM

esop

hyll

um

spec

ies,

mel

ob

esio

ids

ind

et.

>20

mC

ora

l,b

ryzo

ans

A32

Rh

od

oli

thL

ith

oth

amn

ion

spec

ies,

mel

ob

esio

ids

ind

et.

>20

m

A3

isa

lim

em

ud

sto

ne

and

A31

isa

fin

eca

rbo

nat

esa

nd

sto

ne,

bo

thar

eu

nfo

ssil

ifer

ou

s.A

5,A

6,A

19an

dA

30ar

eco

rals

wit

hn

oo

ther

com

po

nen

ts.

Th

eo

ccu

rren

ceo

fco

rall

ines

,u

nle

ssre

-dep

osi

ted

do

wn

slo

pe,

ind

icat

esd

epth

so

f<

120

m.

Bio

stra

tigr

aph

icag

esar

ed

eter

min

edfr

om

larg

efo

ram

inif

era

and

pal

aeo

dep

ths

fro

mco

rall

ine

alga

e,as

dis

cuss

edin

the

text

.

*Scr

ipp

sIn

stit

uti

on

of

Oce

ano

grap

hy

Cru

ise

Ari

esV

IID

red

ge43

.

D. A. Clague et al.

1026 Journal of Biogeography 37, 1022–1033ª 2009 Blackwell Publishing Ltd

The summit consists of a series of terraces varying from

about 60 to 125 m high (Fig. 1d–g), which is a similar height

to that of the series of terraces that surround the northern half

of Hawaii (e.g. Moore & Fornari, 1984; Moore & Clague, 1992)

and Lanai (Campbell, 1986; Faichney et al., 2009).

The present depth of the dredged carbonate samples is

725 ± 100 m. However, the youngest biogenic components

were transported downslope to the location where the

carbonate clasts were deposited, and probably formed on the

large nearly flat summit of the seamount. The present depth of

Table 2 Sr-isotopic data for carbonate

components in K�oko Seamount samples.Sample* Material analysed 87Sr/86Sr* SE (%)

Minimum

age (Ma)

Maximum

age (Ma)

Median

age (Ma)

A5 Coral 0.708034 0.0006 28.36 28.95 28.64

A6 Coral 0.707985 0.0008 29.50 30.35 29.89

A8 Foraminifera/matrix 0.708705 0.0009 16.04 16.46 16.24

A12 Coral 0.708090 0.0008 26.60 27.54 27.07

A18 Coral 0.708079 0.0007 26.99 27.86 27.44

A19 Coralline algae 0.708055 0.0007 27.79 28.45 28.15

A20 Coralline algae 0.708026 0.0008 28.49 29.20 28.84

A22 Coralline algae 0.708093 0.0007 26.53 27.40 26.97

A23 Coralline algae 0.708372 0.0009 20.98 21.42 21.22

A26 Coralline algae 0.708204 0.0008 23.68 24.23 23.96

A30 Coral 0.709168 0.0007 0.08 0.46 0.26

A32 Coralline algae 0.708116 0.0007 25.71 26.71 26.15

*87Sr/86Sr values reported relative to 0.710250 for NBS-987. Ages were determined from87Sr/86Sr using the look-up table version 4: 08/03 after Howarth & McArthur (1997) and

McArthur et al. (2001). The look-up table applies a fifth-order polynomial describing the

relationship between the 87Sr/86Sr of sea water and time. The resulting median, minimum and

maximum age estimates include the uncertainty (2r) in 87Sr/86Sr measurement as well as the

uncertainty (2r) in the calculated mean age from Howarth & McArthur (1997) and McArthur

et al. (2001).

(a) (b)

(c)

(e)

(d)

Figure 2 Photomicrographs of carbonates

from K�oko Seamount. (a) Shallow-water

zooxanthelate coral encrusted by coralline

algae, (b) Amphistegina packstones with

subordinant coralline algae probably formed

at intermediate depths according to the depth

distribution of modern analogues, (c) pack-

stones to floatstones with lepidocyclinids,

bryozoans and coralline red algal nodules

accumulated in deeper-water settings of the

carbonate platform, (d) Lithothamnion is the

main component of the coralline algal

assemblages, (e) Spiroclypeus indicates a

Chattian–Aquitanian age for the deep-water

rhodolith facies. Scale bars are 1 cm in (a)

and (c), and 0.5 mm in (b), (d) and (e).

All samples are from Aries VII Dredge 43.

Geological constraints of Hawaiian lineages

Journal of Biogeography 37, 1022–1033 1027ª 2009 Blackwell Publishing Ltd

this flat summit is between 253 m (the shallowest depth

measured on K�oko) and 343 m (the shallowest depth nearest

the dredge site).

DISCUSSION

By integrating the bathymetry, the ages of formation of the

seamount and of the various facies of carbonate deposits, the

palaeowater depths at which the deposits formed, the latitu-

dinal motion history, sea level and ocean temperature varia-

tions during the history of the seamount, and latitudinal

variations in sea surface temperature, we can reconstruct the

history of subsidence and carbonate bank accumulation on

K�oko Seamount. The history of slow subsidence accompanied

by coral growth that could just keep pace presents the greatest

uncertainty in the discussion to follow.

Bathymetry

The series of terraces on K�oko Seamount appear to be coral

and indicate that the guyot top is covered by a series of coral

reefs that are younger towards the summit, as occurs offshore

Hawaii (Ludwig et al., 1991).

The shallowest point on the summit apparently represents a

small patch reef that continued growing after most of the

carbonate platform had drowned. In our models, we will use

the depth of the larger platform at c. 340 m, the most likely

source of debris sampled in the dredge. The rounded shape of

the clasts supports the interpretation that they have been

transported.

Ages

Sample A23 contains larger foraminifera Lepidocyclina (Eul-

epidina) of Early Oligocene (33.9–28.4 Ma) to Aquitanian (23–

20.4 Ma) age, Spiroclypeus of latest Oligocene (28.4–23 Ma) to

Aquitanian age, and Neorotalia of an Early Oligocene to

earliest Aquitanian age. The Spiroclypeus are early forms

transitional to Vlerkina, indicating the young end of the range,

namely an Aquitanian age (Lunt & Adams, 2004; Renema,

2007). Sample A8 contains Amphistegina, Operculinella and

Operculina. The frequent presence of Operculinella indicates a

Burdigalian (20.4–16 Ma) or younger age (Renema, 2007).

This sample also has an age based on Sr-isotopes of

16.15 ± 0.6 Ma, consistent with the Burdigalian foraminiferal

age.

The Sr-isotope-derived ages of 27.1 ± 0.5 to 29.9 ± 0.4 Ma

for four of the five corals and an algal coating on another coral

may represent the youngest corals on K�oko Seamount. We

interpret the single Pleistocene age to be a recrystallization age.

The oldest corals presumably began growth soon after the

volcano formed at c. 50.4 ± 0.1 Ma, during the Ypresian stage

of the Eocene (55.8–48.6 Ma, all ages based on International

Commission on Stratigraphy, 2004). These ages suggest that

K�oko Seamount was probably capped by an active shallow-

water coral reef for about 23 Myr between 50 and 27 Ma.

Ages of rhodoliths, also based on Sr-isotopes, range from

28.8 ± 0.4 to 21.2 ± 0.2 Ma, and a single sample of Amph-

istegina-rich sediment from a rhodolith facies rudstone yields

an age of 16.2 ± 0.5 Ma. The rhodolith ages overlap with the

coral ages and suggest simultaneous deposition of the two

facies at different depths, with the corals at < 20 m and the

rhodoliths downslope at < 120 m.

The palaeontological ages are within the range of Late

Oligocene or Early Miocene reported from the small sample of

drill cutting recovered at DSDP site 309, but younger than the

late Eocene (37.2–33.9 Ma) shallow-water large foraminiferal

ages reported from sites 308 and 1206. Carbonate deposition

began shortly after the volcano was constructed at 50 Ma

(Sharp & Clague, 2006) and continued in both shallow- and

deep-water settings across the bank, at least intermittently,

until c. 16.2 ± 0.5 Ma, when the relatively deep-water carbon-

ate deposition ceased.

Palaeoenvironments

Most zooxanthelate corals live in the upper 20 m (Grigg &

Epp, 1989), although they can survive to depths as great as

90 m, where they exhibit very low growth rates. The samples

do not allow us to assess whether they formed any kind of reef,

although the seismic reflection data (Greene et al., 1978, 1980)

suggest that significant, perhaps reefal, carbonate deposits exist

on the summit of K�oko Seamount. The presence of ooids near

the bottom of the sediment section above basement rocks from

site 308 (Matter & Gardner, 1975) is consistent with warm,

shallow conditions (Simone, 1981). The foraminifer Amph-

istegina, a common component in the bioclastic packstone,

rudstone and rhodolith facies, lives to depths of 100 m in

present-day Pacific benthic settings (e.g. Hohenegger, 2004;

Renema, 2006), but its depth distribution is much reduced in

north Pacific settings and it is restricted to c. 30 water depth

in Hawaii (Hallock, 1984). The thick-biconvex shape of the

Amphistegina tests in many of the K�oko samples suggests

depths shallower than 50 m (Hallock, 1987). However, down-

slope displacement of foraminiferal tests during packstone–

rudstone accumulation cannot be discounted. The presence of

coralline assemblages that are composed only of melobesioids

suggests palaeodepths of more than 20 m (Adey et al., 1982).

The abundance of bryozoans and the scarcity of corals in these

samples suggest deposition in cooler or deeper waters than the

coral facies (Schlanger & Konishi, 1975).

Coralline algal assemblages dominated by Lithothamnion

and Sporolithon occur in the Hawaiian Islands on seafloor

deeper than 50–60 m (Adey et al., 1982). The maximum

growth depth for these rhodoliths is estimated at 100–120 m,

as below these depths coralline red algae growth is limited to

very thin crusts, if it occurs at all (Braga et al., 2005).

The simplest interpretation of the carbonate facies recorded

on the top of K�oko Seamount is that they reflect a progressive

drowning of a carbonate platform, completed near the end

of the Burdigalian stage of the Early Miocene. The method of

sampling prevents any depth or stratigraphic arrangement

D. A. Clague et al.

1028 Journal of Biogeography 37, 1022–1033ª 2009 Blackwell Publishing Ltd

of facies, but the regional context of seamount evolution allows

the interpretation that zooxanthelate corals grew on top of the

seamount soon after the volcano formed and, owing to the

continued subsidence of the seamount, were later overlaid by

bioclastic sediment rich in Amphistegina, formed at similar to

greater depths. The final, youngest deposits recorded on the

seamount are of the facies that forms in the deepest water, the

rhodolith floatstones, which accumulated on the summit of the

seamount at 60–120 m depth, before their demise in the Early

Miocene.

Subsidence

The Emperor Seamounts, like their Hawaiian Island progeny,

underwent two distinct stages of subsidence. As the volcano

was actively growing, and continuing for about 1 Myr after

eruptions became infrequent, there was a period of rapid

subsidence caused by the flexure of the lithosphere as it

isostatically adjusted to the added weight of the volcano

(summary in Moore, 1987). Once equilibrium had been

achieved, the volcanoes slowly continued to subside as the

underlying plate aged, cooled and contracted. The K�oko data

allow us to assess both of these periods of subsidence.

Using the present depth at which the carbonates probably

formed (c. 340 m), the age (27.1 Ma) and palaeodepth (20 m)

for the corals, and the age (16.2 Ma) and depth (120 m) of the

last deep-water carbonate deposition, we can calculate that the

subsidence rates of K�oko Seamount were: (1) c. 0.009 mm yr)1

from 27.1 to 16.2 Ma, (2) c. 0.014 mm yr)1 from 16.2 Ma to

present, and (3) c. 0.012 mm yr)1 over the entire last

27.1 Myr. These rates are all statistically the same and indicate

an average subsidence rate of 0.012 ± 0.003 mm yr)1. This

very slow calculated subsidence rate is consistent with rates

deduced from the thermal cooling and contraction of an old

ocean plate (Stein & Stein, 1992). At this rate, K�oko Seamount

would have subsided an average of 590 m since the volcano

attained isostatic equilibrium at c. 49 Ma.

If we subtract the post-isostatic subsidence of 590 m from

the 1555 m of total subsidence indicated by the depth of the

break-in-slope, we are left with 965 m of subsidence between

the end of volcano construction and the attainment of isostatic

equilibrium. The ocean crust beneath K�oko was younger than

that beneath Hawaii (60 vs. 90 Ma) when the volcanoes

formed (Clague & Dalrymple, 1987), so the lithosphere should

have been less rigid and the flexure resulting from loading

somewhat greater, so one could argue that the depth to the

break-in-slope should be somewhat greater than the 1000–

1100 m observed around the main Hawaiian Islands. On the

other hand, this greater flexure should also have occurred

earlier in the history of the volcano, as a weaker lithosphere

would equilibrate under the growing load of the volcano more

quickly, which might make the break-in-slope shallower than

observed around Hawaii, and consistent with our calculations.

This uncertainty in the earliest subsidence history of the

volcano and the unknown growth rate of the coral reef as a

function of time between 49 and 27 Ma obscure our under-

standing of the early subsidence history of K�oko Seamount.

The unknown growth rate of the reef will also be a factor when

we try to determine when the summit of K�oko Seamount

submerged.

Latitudinal migration of K�oko Seamount

Hotspot models (Wilson, 1963; summarized in Clague &

Dalrymple, 1987) predict that the volcanoes of the Emperor

Seamounts are being carried passively on the tectonic plate

after forming over the plume, which is now under Hawaii.

Greene et al. (1978) first used reefs on the Emperor Seamounts

to argue for northward movement of the volcanoes since their

formation. Palaeomagnetic data (summarized in Clague &

Dalrymple, 1987), confirmed by more recent data (Tarduno

et al., 2003), show that the hotspot has not always been fixed,

at least during the formation of the Emperor Seamounts. In

the first 5 Myr of its life, K�oko Seamount migrated northwards

as the hotspot migrated southwards (Tarduno et al., 2003).

During this period, the seamount moved northwards with the

plate at about 69 km Myr)1 from c. 21.5� N to c. 23� N. It

then migrated northwards at about 31 km Myr)1 from 45 Ma

until present as the Pacific plate moved west-north-west to

reach its present latitude of c. 35� N. When the youngest corals

died out at 27 Ma, K�oko Seamount was located at 27.8� N, just

south of present-day Midway Atoll (Fig. 1b). When deep-

water carbonate deposition ceased at 16.2 Ma, K�oko Seamount

was located at 30.8� N, near present-day Hancock Seamount

(Fig. 1b), nearly 450 km north-west of Midway Atoll.

Climatic conditions as K�oko Seamount subsided and

migrated northwards

At 50.4 Ma, when K�oko Seamount formed at 21.5� N,

conditions should have been tropical and coral growth rates

high because it was about the time of the Early Eocene Climatic

Optimum (Zachos et al., 2001). The presence of ooids in the

Site 309 samples (Matter & Gardner, 1975) is consistent with

this warm, shallow interpretation of conditions soon after the

seamount formed. The summit of K�oko Seamount was

subaerial at this time. Corals should have produced fringing

reefs that migrated upslope as the island continued to subside,

similar to those that surround Hawaii today (e.g. Webster

et al., 2007). The single swathe of multibeam data (Fig. 1)

supports this concept as it shows numerous terraces with steep

faces, ranging in height from c. 60 to 150 m. For the next

16 Myr, global temperatures steadily decreased, and by 34 Ma

bottom water temperatures were nearly 7 �C cooler than at the

Early Eocene Climatic Optimum (Zachos et al., 2001).

At 34 Ma, Antarctic ice sheets (the Oi-1 glaciation) began to

form for the first time in the Cenozoic, and sea level dropped

by about 55 m (Miller et al., 2008) to 70 m (Coxall et al.,

2005) as bottom ocean temperatures dropped by another

2–3 �C (Zachos et al., 2001). Following the Oi-1 glaciation,

temperatures remained cool until the Late Oligocene Warm-

ing, which began about 26 Ma. Near the end of this cool

Geological constraints of Hawaiian lineages

Journal of Biogeography 37, 1022–1033 1029ª 2009 Blackwell Publishing Ltd

period the last shallow-water corals on K�oko Seamount

apparently died. Temperatures then remained generally warm,

with a brief glacial (the Mi-1 glaciation at about 24 Ma) until

after the youngest age (16.2 Ma) of the carbonate samples

recovered from K�oko Seamount.

Submergence of the top of K�oko Seamount

Any vestige of K�oko Island subsided below sea level sometime

before the final coral died out at 27.1 Ma. Estimating when this

occurred requires data on the growth rate of the corals and the

subsidence rate at the time. Using our estimated subsidence

rate of 0.012 mm yr)1 implies submergence of the coral cap by

c. 29 Ma, or somewhat earlier if the corals were partially

keeping up with subsidence. We suggest that the summit might

have submerged prior to 34 Ma, but then re-emerged when sea

level fell by 55–70 m (Coxall et al., 2005; Miller et al., 2008)

during the Oi-1 glaciation (Zachos et al., 2001), only to

resubmerge as sea level rose 1–1.5 Myr later, presumably at a

rate faster than the growth rate of the corals under such cool

conditions. The summit of K�oko Seamount probably sub-

merged when sea level rose rapidly at 32.5–33 Ma (Haq et al.,

1987; Miller et al., 2008), in reasonable agreement with the

simple subsidence model calculations of Clague (1996) of

36.5 Ma. This scenario would imply that corals were able to

grow at rates of about two-thirds of the subsidence rate, even

at the northerly latitude of the seamount and given the cool

climate conditions at that time.

The estimated 16.5–17 Myr life span of K�oko Seamount as a

rocky island is longer than the ages of the smallest and oldest

volcanic island remnants in the Hawaiian chain. Gardner and

La Perouse Pinnacles have interpolated ages of 15.8 and

12.9 Ma (Clague, 1996) and poorly constrained K–Ar ages of

12.3 and 12.0 Ma, respectively.

Why did the reef and then the deep-water rhodoliths

stop growing?

The carbonate bank or reef on K�oko Seamount was confronted

by a confluence of adverse trends after reef growth began

50 Ma. The seamount was moving northwards into cooler

waters as global climate was cooling (bottom waters had

cooled by 7 �C by the time of the Oi-1 glaciation at 34 Ma),

and the island subsided c. 1100 m in the first 1 Myr, and

continued to subside slowly after that.

It is difficult to explain the drowning of shallow-water

zooxanthellate coral deposits and the subsequent deposition of

deeper facies (bioclastic packstones and rhodolith floatstones)

by subsidence alone, once the initial isostatic phase of rapid

subsidence was completed. The long-term subsidence rate is

far lower than the accretion rate of shallow coral reefs

(Buddemeier & Smith, 1988; Smith & Buddemeier, 1992;

Webster et al., 2007). K�oko Seamount simply migrated north

away from a contracting belt of tropical waters where coral

reef growth took place in the Central Pacific. We estimate

that the last of the corals gave up about 27.1 Ma, having built

the estimated 530-m thick carbonate deposits on top of the

seamount during a 23-Myr period after the volcano formed.

The coral cap apparently survived, or was revived by, the Oi-1

glaciation at 34 Ma, but had died out and submerged by the

time the Late Oligocene Warming began c. 26 Ma (Zachos

et al., 2001).

After the summit had submerged below 20 m depth at

c. 27.1 Ma, the deep-water carbonate factory was established

and lasted until the Burdigalian stage of the Early Miocene,

when the combination of very low accretion and growth rates

and slow subsidence brought the summit below the deep-water

coralline algal factory. Sea level rise associated with warming at

the end of the Oligocene (Miller et al., 2008) and in the Early

Miocene (Haq et al., 1987; Abreu et al., 1998) may have

assisted in their drowning, but continued slow subsidence and

even slower deep-water coralline growth probably account for

their eventual death.

Implications for Hawaiian biogeography

The submergence of K�oko Seamount at c. 33 Ma supports the

model proposed by Clague (1996) that successive colonization

of the Emperor and Hawaiian Islands (island hopping) ceased

during the Late Eocene to Early Oligocene. K�oko Seamount is

the key to understanding the colonization of the Hawaiian

Island chain because it was so large and survived for so long as

an island. No other island was comparable in size until

Gardner Pinnacles formed c. 16 Ma. Between c. 33 Ma, when

K�oko Seamount submerged, and 30 Ma, when Kure Island

grew, there were only transient islands (lasting < 0.5 Myr) in

the Hawaiian chain (Fig. 1b). K�oko Seamount was quite

distant from these ephemeral islands and was itself a flat,

low-lying coral island with limited terrestrial biodiversity. Kure

Island and Pearl and Hermes Reef each were islands that

reached elevations of perhaps 870 m (Clague, 1996), but they

were small, low, distant islands by the time Lisianski Island, the

next high island (> 1000 m high) formed about 23 Ma. Until

that time, individual small islands existed only for brief

periods and submerged quickly. Between the submergence of

K�oko Seamount at 33 Ma (or even slightly earlier) and the

formation of Lisianski Island at 23 Ma, the transmission of

local terrestrial and shallow marine propagules from island to

island along the chain was probably limited to lowland species.

Islands of the chain starting with Lisianski had to be colonized

by migrants from distant shores, in broad agreement with

MRCA analyses of Hawaiian lineages.

CONCLUSIONS

The 50-Myr-old K�oko Seamount initially had a substantial

shallow-water carbonate reef build-up that contained zooxan-

thelate corals. These shallow, warm-to-temperate-water reefs

came under increasing stress over the next 23 Myr as global

climate cooled and the seamount migrated northwards and

subsided slowly (c. 0.008 mm yr)1) as a result of the cooling of

the aging underlying lithosphere. The summit submerged

D. A. Clague et al.

1030 Journal of Biogeography 37, 1022–1033ª 2009 Blackwell Publishing Ltd

c. 33 Ma, slightly after the first glacial period of the Cenozoic,

and the coral reef probably died as a result of unsuitable water

temperatures by 27.1 Ma. The volcano continued its slow

subsidence and first accumulated shallow-to-mid-depth fora-

minifer Amphistegina in bioclastic packstones and then deep-

water rhodoliths, until the end of the Burdigalian stage of the

Early Miocene (16 Ma), when the deep-water facies died out as

a result of the combination of slow subsidence rates and even

slower deep-water carbonate accumulation rates.

Carson & Clague (1995) and Clague (1996) proposed that

biota on older Hawaiian–Emperor Islands could not island-

hop along the entire chain to eventually colonize the present

Hawaiian Islands. The new data presented here from K�oko

Seamount provide independent evidence confirming the

model proposed by Clague (1996). There was a time period

from about 33–30 Ma in which no high islands, and only

transient low islands, existed. All terrestrial life-forms from the

older portion of the chain would have been extinguished. Kure

Island and Pearl and Hermes Reef formed moderate-sized

islands (c. 870 m tall) 3–7 Myr after K�oko Seamount

submerged, but they were small, low distant islands by the

time the next large islands began to grow, starting with

Lisianski Island c. 23 Ma. The ancestors of the modern

Hawaiian terrestrial and shallow marine biota were unlikely

to have become established before 23 Ma, when distant

colonizers, perhaps supplemented by colonizers from low

distant islands such as Kure Island and Pearl and Hermes Reef

(250–500 km away), could arrive, radiate and migrate from

island to island until some of them reached the present islands.

ACKNOWLEDGEMENTS

We thank Warren Smith at the Scripps Institution of Oceano-

graphy Core and Dredge Sample Repository for locating the

samples, and the Museum of Paleontology at the University of

California, Berkeley, for their unsuccessful attempt to locate the

corals examined by Wyatt Durham in the early 1970s. We also

thank Jennifer Paduan for making the maps.

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BIOSKETCH

David A. Clague is a marine geologist specializing in

volcanoes in the oceans. He has studied the volcanic formation

of the Hawaiian Islands and Hawaiian–Emperor volcanic

chain, their destruction through subsidence and giant land-

slides, and the formation of coral reefs on the rapidly

subsiding present-day islands. He is a co-editor of a mono-

graph on Explosive Subaequeous Volcanism and of the newly

released Encyclopedia of Islands (University of California

Press). The Encyclopedia examines the role that geological

history plays in the biodiversity dynamics on islands.

Author contributions: D.A.C. conceived the study and its

scope, located the samples, did the subsidence calculations,

and wrote most of the manuscript; J.C.B. identified the

corallines and wrote part of the manuscript; D.B. and W.R.

identified the large foraminifera and determined their pala-

eontologic ages; P.D.F. performed the Sr-isotopic analyses;

J.M.W. developed the study concept and methodology with

D.A.C. and refined the paper.

Editor: Kostas Triantis

This paper is an additional contribution to the Special Issue

that arose from the symposium Evolutionary islands: 150 years

after Darwin (http://science.naturalis.nl/darwin2009), held

from 11 to 13 February 2009 at the Museum Naturalis,

Leiden, The Netherlands. The theme of the symposium was to

explore the contribution of islands to our understanding of

evolutionary biology and to analyse the role of island biological

processes in a world in which the insularity of island and

mainland ecosystems is being drastically altered.

Geological constraints of Hawaiian lineages

Journal of Biogeography 37, 1022–1033 1033ª 2009 Blackwell Publishing Ltd