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Biological Journal of the Linnean Society, 8: 2 0 5 - 2 1 6 . With 3 figures
September 1 9 7 6
A geological perspective of the upland biota of
Laysan atoll (Hawaiian Islands)
S. 0. SCHLANGER
Geological Institutes, Universities o f Leiden and Utrecht, The Netherlands
AND
G. W. G I L L E l T F.L.S.t
Department oj’Bio1og.v. University of California, Riverside
Accepted for publicution November 1 9 7 5
Laysan Island, a low atoll in the Leeward Hawaiian Islands, exhibits a biota that contains nine
genera of upland and montane lineages of animals and plants usually of restricted range and
occurrence and adapted t o high island habitats. The geological history of Laysan Island,
interpreted in terms of plate tectonic theory and recent ideas on the formation of linear island
chains, shows that Laysan existed as an active high volcanic island approximately 1 5 million
years ago. Since that time subaerial erosion and tectonic subsidence have combined to reduce
the height of the island. During the past 2 5 0 , 0 0 0 years glacio-eustatic sea level changes have
resulted in Laysan Island fluctuating, geomorphologically, between a high limestone island and
an atoll status. Laysan is viewed as a refugium for upland and montane lineages able to keep
pace, via great adaptive flexibility, with drastic habitat changes. The thesis that plants of Laysan
are recent arrivals is considered unlikely, as regards the upland contingent, in view of the
geological history of the island.
CONTENTS
Introduction
. .
Discussion
. .
Conclusions
. .
Acknowledgements
References
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205
207
213
214
214
IN’IKODUCIION
Biological surveys of the Leeward Hawaiian Islands, carried out over the past
1 0 0 years (Rothschild, 1893-1900; Bitter, 1900; Christophersen & Caurn,
1931; Lamoureux, 1963; Ely & Clapp, 1973) show the presence on Laysan
t Deceased.
205
206
S. 0. SCHLANCER AND G . W . GILLE'M
Table 1. Occurrence in the Hawaiian Chain of land animals and upland
angiosperms of Laysan and Nihoa
Laysan
Nihoa
Kauai
Oahu
Molokai
Maui
Hawaii
X*
X
X
X
X
X
X
Land snails ( 1 )
Tornatellides
Land birds
Rails (Rallidae) ( 2 )
Porzana
X*
X
X
tioneycreepers (Drepanidae) ( 3 )
Hima tione
Psittirostra
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X*
X
X
X
X*
X
X
X
X
X
X
X*
X
X
X*
X
Warblers (Sylviidae) ( 3 )
Acrocephalus
Upland angiosperms
Sandalwood (Santalaceae)
Santalum (4)
X*
Pinks (Caryophyllaceae)
Schiedea ( 5 , 6 )
Mint (Lamiaceae)
Phyllostegia (7.8)
Aster (Asteraceae)
Lipochoete ( 9 )
Palm (Arecaceae)
Pritchardio (1 0 )
X
X*
X
1 . Cooke and Kondo, 1960. 2. Olson, 1973. 3 . Berger, 1972. 4. Rock, 1916. 5. Hillebrand, 1888
6. Herbst, 1975. 7. Bitter, 1900. 8. Sherff, 193Sb. 9 . Sherff, 1935a. 10. Beccari & Rock, 1921.
= extinct
Island, now a low atoll, of both animals and plants representative of upland and
montane lineages. I t must be emphasized that the insular biotas of the Pacific
include two broad habitat groups: (1) upland and montane animals (snails,
insects, birds, bats) and plants often of restricted range and occurrence, both
adapted to habitats above the strand, on high islands; and, (2) strand animals
and plants of widespread occurrence on atolls and on the beaches of high
islands. The strand plants are halophytes with a tolerance of coral, saline soils,
salt spray, and other rigours of the beach habitat. T h e biota of Laysan, the
youngest atoll of the Leeward group, include a generous representation of
marine animals, particularly marine birds, and strand plants of this second
group. I t is most significant, however, that the Laysan biota also include nine
genera of upland and montane lineages of animals and plants characterized in
the first group. These are listed in Table 1. The table excludes Arthropoda even
though Nihoa and Laysan almost certainly have had arthropods of similar
habitat relationships (Carson, Hardy, Spieth & Stone, 1970).
The history of cxploration and of the impact of Man on Laysan Island is
reviewed by Rothschild (1893-1900), Bitter, (1900), Christophersen & Caum
(1931), and Ely & Clapp (1973). Accounts begin with the observations of
Isenbeck, surgeon of the vessel Moller, Capt. Stanikowitch, in 1828. These and
later records portray a saga of the destruction and extinction of a rare, upland
biota of which only one element, the Laysan finch (Psittarostra),of Group 1
survives. Infrequent reports between 1828 and 1890 give accounts of an
extensive vegetation cover composed of grasses, low shrubs (shrubs of Santaltim
up to 2.5 m high and 10-15 cm diameter), and of a small population of the fan
LAYSAN ATOLL
207
palm, Pritcliartlia (five or six trees reported in 1857). Fresh-water was obtained
from wells. A peat deposit was reported (Schauinsland, 1899), a unique
attribute for an atoll of perhaps 1 0 m elevation. This very confined and
apparently fragile, upland ecosystem was subjected to violent disruption with
the onset, in about 1890, of guano surface mining on the south portion of the
island, In June, 1891, Palmer and Munro, collectors for Rothschild, visited
Laysan and reported only dead remnants of f’ritchurtlia. apparently the first
upland element to fall to extinction. Schauinsland visited the island in 1896
and made the first known collections of plants, these named and reported by
Bitter (1900). Listed by Bitter were 26 species of vascular plants, these with
Pritcliardia totalling 27 species (Lamoureux, 1963), the only available
compilation of the pristine flora of Laysan. Four of these (I’ritcliartliu,
Sutitaliirn. Plijdlostc~giu,and Lipocliactu) o r approximately 15?6 of the flora are
here classified as an upland, non-strand contingent. By 1910 the guano
operations had become defunct, but in about 1903 rabbits and guinea-pigs were
released on the island, then under private ownership. These accomplished a
swift destruction of the indigenous vegetation and the demise of upland bird
populations which earlier had been estimated at hundreds of individuals. The
slaughter of hundreds of thousands of marine and land birds by feather
poachers in 1909 and 1910 contributed to the scene. By 1923 Satitalirtn,
Lipocliaeta, and Pli.i*llostegia were extinct and the island was nearly bare of
vegetation beyond the shore line. T h e Laysan Millerbird (Acroceplialzrs) and
the Laysan Honeycreeper (Himiatone) had also met extinction by 1923. The
Laysan Rail (Porzaria) persisted until its extinction on Laysan in about 1936. I t
had been released on Midway, but became extinct there in 1943. By 1923,
when the rabbits and guinea-pigs were purposefully eliminated from Laysan, it
could have been regarded as a ravaged, desert island, the destruction having run
its full course. Laysan’s present biota, expressed in terms of indigenous
elements, is a strand and marine biota with the sole exception of the Laysan
finch (Psittarostra), whose survival undoubtedly may be assigned to its
omnivorous diet, comprised largely of the eggs of marine birds.
In recent years the geological history of the Hawaiian Islands has been
interpreted within the framework of new models postulated to account for the
formation of the ocean floors and the formation of linear island chains in
general (Wilson, 1963; Morgan, 1968, 1971, 1972; Jackson, Silver &
Dalrymple, 1972; Jackson, in press). I t is the purpose of this paper to relate the
presence of upland and montane animal plant lineages on Laysan Island to the
geological history of Laysan as it evolved from a high volcanic island to a low
atoll.
DISCUSSION
The Hawaiian Islands, like so many other island groups in the Pacific Basin,
are not simply clustered about some point b u t are arranged in a definite, linear
chain. The Hawaiian Island chain extends from the island of Hawaii to Midway
and Kure atolls (Fig. 1). Over 100 years ago Dana (1849) observed a regular
geomorphic gradient along the Hawaiian chain. The active, growing volcanic
islands, such as Hawaii, are restricted to the southeast end of the chain. Extinct
volcanic islands such as Molokai, Oahu, Kauai, and Niihau to the northwest of
S. 0. SCHLANGEK AND C. W . CILLETT
208
60'
500
40'
30°
200
OE
I
/70°
I
/BOO
I
I
/ 70'
/60°
1
15 ' W
Figure 1. Map of the Hawaiian Island Chain and its submarine extension to the northwest--the
Emperor Seamount Chain. The arrows and the ages show the suggested direction of Pacific
Plate motion during the indicated periods (based o n data from Morgan, 1972).
Hawaii show a degree of erosion proportional to their distance from Hawaii.
Farther still to the northwest are the islands of Nihoa, Necker, and Gardner
Pinnacles-now mere remnants of former large volcanoes, as shown by the
bathymetry of their immediate surroundings, that have been eroded almost
down to sea level and encircled by coral reefs (Fig. 2). At the northwest end of
the Chain are the atolls of Midway and Kure, where n o volcanic rock is found
at sea level. Dana drew the inference that each island in such a chain was at a
particular stage in the life history of an atoll as earlier postulated by Darwin
(1842).Darwin's subsidence theory held that an atoll began as a high volcanic
island which subsided as volcanism ceased and the sea floor it rested upon
deepened. As long as the island remained in a tropical environment where reef
growth could keep pace with subsidence, an atoll developed on the sinking
volcanic foundation. His subsidence theory was upheld by results of drilling at
Eniwetok (Schlanger, 1963). Recent studies on the eruption rate of Kilauea
volcano on Hawaii and estimates of the volume of the volcanic edifice that
forms the island show that the entire structure could have been built in a few
hundreds of thousands of years; the calculated rate of eruption of Kilauea is
0.11 km3/year (Shaw, 1973) and the volume of the structure is 19,400km3
(Bargar & Jackson, 1974).According t o Dana's view (1849)the islands along
LAYSAN ATOLL
E'OTO'I
t'O~0'01
C 'OTL' I I
333Y 0 8 V W
NVSAVl
NNVB Y 3 3 N O l d
IYSNVISIl
209
2 10
s. 0 . SCIH.ANGI<K AND
G.
w. c i i . m T r
the Hawaiian Chain should show an increase in age, in terms of the date of
cessation of volcanism, from the presently active island of Hawaii to the long
extinct volcanic island below Midway Atoll. Drilling at Midway (Ladd, c't ul.,
1970) confirmed his inference. The volcanic basalt basement, dated a t
17.9 ? 0.6 million years BP (Dalrymple, Lamphere & Jackson, 1974) was
reached below 385 m of coral limestone that accumulated as the cooling
volcanic foundation subsided. Since 1964 (McDougall, 1964; McDougall &
Swanson, 1 9 7 2 ; Doell & Dalrymple, 1973; Dalrymple et ul., 1974; Clague,
Dalrymple & Moberly, in press) the dates at which major volcanism along the
chain essentially ceased have been determined in some detail (Fig. 2).
The rate a t which the height of the volcanic edifice above sea level is reduced
with time is shown by the dashed line on Fig. 2, which was fitted by eye to the
present elevation of volcanic rocks along the chain from the island of Hawaii to
the basalt basement reached by the drill below Midway Atoll. In the Hawaiian
chain an island appears to progress in a general way from the active volcanic
stage to the atoll stage in approximately 15 million years. The dashed line on
this figure is constructed on the argument that: (a) Laysan Island and Mar0
Reef, which are probably on top of the same large volcanic edifice, are the
most south-easterly islands that appear, on a geomorphic basis, to have very
recently become atolls; and (b) that the date at which volcanism ceased there
was approximately 15 million years BP, a date that can be derived by
interpolating between Midway Atoll, where volcanism ceased approximately 18
million years BP and La Perouse Pinnacle where volcanism ceased approximately 1 2 million years BP (Fig. 2).
In addition to t h e vertical component of island history, each island in the
Hawaiian Chain has a history of horizontal motion because the volcanic edifices
that form the foundations below the islands have formed through the
accumulation of lavas on top of the moving Pacific Plate. Acording to the
plate tectonic model (Le Pichon, 1968; Morgan, 1968; Isaks, Oliver & Sykes,
1968; Heirtzler et ul., 1968) the surface of the earth can be divided into a
relatively small number of lithospheric plates, perhaps 100 km thick, that move
over the asthenosphere. The Pacific Plate, upon which the Hawaiian Island
chain rests, is bounded on the west by the Marianas-Japan-Kurile
trenches
and on the east by the East Pacific Rise that extends from Central America to
between Antarctica and Australia. The plate is moving in a northwesterly
direction (Winterer, 1973; van Andel, 1974) a t the present and has been
moving in this direction for the past 4 2 million years. Prior to that time the
Pacific Plate probably moved in a more northerly direction (Morgan, 1972;
Jackson et al., 1972; Clague & Jarrard, 1973) for some tens of millions of years
(Fig. 1). These directions of motion have been deduced from data such as the
azimuthal trends of the major linear island chains in the Pacific, seismic
first-motion studies, and analyses of the paleolatitude position of seamounts,
within which are preserved the records of ancient magnetic fields (Francheteau,
Harrison, Sclater & Richards, 1970). Paleomagnetic studies of basalts cored
below Midway Atoll show that the volcanic basement there could have formed
at a latitude of 15" + 4.0'-a latitude consistent with their possible origin at the
present latitude of Hawaii (Grommk & Vine, 1972).
The mechanism by which lava is generated below the lithospheric plate and
LAYSAN ATOLL
211
breaks through the plate to form volcanoes is a subject of intense debate and
will not be treated here.
If an observer could have spent the past 70 million years looking down on
the Pacific Ocean watching the evolution of the Hawaiian and Emperor Chains
(i.e., taking the bird’s eye view, which is most germane to this paper) each
island in the Hawaiian Island Chain and now the completely submarine
Emperor Seamount Chain, would have exhibited the following history. Each
island would have been seen emerging from the sea as an active volcano near
the present position of the island of Hawaii. This new island would have begun
moving to the northwest as volcanism ceased and the island was carried on top
of the Pacific Plate to the northwest along the path shown by the arrows in
Fig. 1 . Upon cessation of major volcanism, the date of which is shown for each
island where data are available (Fig. 2), the high volcanic island would have
been rapidly reduced by erosion and tectonic subsidence to a relict volcanic
peak with a fringing coral reef. By approximately 15 million years after the
cessation of volcanism the former island would have become an atoll. As
subsidence continued and as these new atolls also continued moving to the
northwest, out of the tropical latitudes hospitable to reef growth, even these
reefs would drown and disappear from view as did Koko Seamount, which
bears on its submerged top, a drowned coral limestone cap which evidently
passed below the coral reef building depth approximately 9 million years BP
(Davies, White & Clague, 1972; Larson et al., 1974).
The geological history of any island in the Hawaiian chain can be
reconstructed within the framework of the general model discussed above.
Further, by considering the effect on island geomorphology of known
glacio-eustatic sea level changes, selected stages in the island’s history can be
examined in detail. Figure 3 is a schematic reconstruction of the geological and
geomorphic history of Laysan Island. The volcanic edifice that must lie below
the island has not been dated, but by interpolating between La Perouse Pinnacle,
dated at 11.7 + 0.4 million years BP and Midway Island, dated at 17.9 + 0.6
million years BP, the cessation of volcanism at Laysan Island between 14 and 15
million years BP can be assumed. At an average eruption rate of 0.034 km3/y
of rock (Shaw, 1973) the Laysan Island-Mar0 Reef edifice, approximately
36,500 km3 in volume (Bargar & Jackson, 1974), could have formed in
approximately 1 million years. Thus, between about 1 5 and 16 million years
BP volcanism at the site of Laysan Island produced a high volcanic island
(Fig. 3A). The cross-sectional area of the Laysan Island-Mar0 Reef edifice
(Fig. 2) indicates that when volcanism ceased the proto-islands of Laysan and
Mar0 probably resembled the Molokai-Maui edifice in terms of both area
exposed above sea level and elevation. Between 1 5 and 9 to 10 million years RP
the height of the island was drastically reduced due to subaerial erosion and
tectonic subsidence. A glacio-eustatic drop in sea level that took place during
this time interval caused the emergence of Eniwetok, Bikini, and Midway atolls
(Schlanger, 1963; Ladd, Tracey & Gross, 1970; Schlanger & Douglas, 1974),
converting them into high limestone islands; presumably proto-Laysan Island
also underwent emergence. By 9 million years BP these high limestone islands
had submerged and become atolls once again (Fig. 3B). Proto-Laysan Island
had by then acquired a mature subaerial profile and a fringing coral reef.
212
S. 0. SCHLANCER AND G. W. CILLETT
1
I
I
YEARS, B.P.
Figure 3. Geological and geomorphic history of Laysan Island. The horizontal scale in years BP
is logarithmic. Diagrams A to G represent the geology of Laysan Island at the ages indicated.
The numbers in parentheses (- 26" N) represent paleolatidudes inferred from the paleolatitude
determined for the volcanic basement below Midway Atoll (CrommC & Vine, 1972). The
sea-level fluctuation curve (inset H) is based on data from Chappel (1974).
Between 9 and 10 and 2 and 3 million years BP erosion of the rapidly
diminishing subaerial volcanic mass continued ; twice more sea level dropped,
during the Messinian Regression 5 to 6 million years BP and during an
Antarctic glacial advance between 3.7 and 2.7 million years BP (Ciaranfi &
Cita, 1973; Cita & Ryan, 1973; Ciesielski & Weaver, 1974). These sea level
fluctuations allowed the development of complex limestone rims around the
deeply eroded volcanic cores (Fig. 3C,D). Laysan during sea level lowerings,
was a Makatea-Henderson type island and resembled presentday Mangaia and
Atiu Islands in the Cook Chain (Marshall, 1927, 1930). Makatea is
characterized by a raised, annular rim of hard limestone, the former encircling
reef, that surrounds a low-lying, central core of volcanic rock-the former
lagoons are now valleys separating the core and rim. During the past .25 my of
its history (Fig. 3E,F,G) the geomorphology of Laysan was largely controlled
by eustatic sea level changes (Chappel, 1974); tectonic subsidence over this
period probably amounted to about 1 5 m. During low stands of the sea such as
took place 18,000 BP Laysan was a high limestone island with perhaps wen the
inner volcanic core covered by limestone. As sea level rose to its present
position, Laysan was rapidly converted into its present form.
In summary, the amount and type of geomorphic change per unit time
exhibited by an island in a chain such as the Hawaiian Chain, varies according
LAYSAN ATOLL
213
to its absolute age. In its youth an island changes from a high, volcanically
active island into a low-profile, extinct volcanic island in a few millions of
years. In its old age an island can change from a high limestone island into an
atoll and vice versa in a few thousands of years due to glacioeustatic sea level
changes.
CONCLUSIONS
In light of the above discussion, Laysan is probably best viewed as a
refugium for an assemblage of upland lineages that were able to keep pace,
through great adaptive flexibility, with the drastic habitat changes imposed by
tectonic subsidence, erosion, and Pleistocene fluctuations in sea level. The most
recent, impressive of these probably was a dtnouement of the rise in sea level
concurrent with the melting of the Pleistocene ice caps, producing a
constriction of the upland habitat and truncating the ranges of its biota. The
survival on Laysan of upland lineages of animals and plants is correlated with
the relatively large surface of the island (c. 300 ha, greater than the combined
areas of the other Leeward Islands).
One of the most impressive qualities of the Hawaiian biota is the propensity
for evolutionary change expressed by broad and abundant adaptive radiations
in many genera. Some plant genera, including Bidens (Asteraceae), Scaevola
(Goodeniaceae), and M y o p o m m (Myoporaceae) have produced ecotypic
populations that occupy diverse habitats from over 1500 m to within 10 m of
the sea. This adaptive diversity has been expressed by all of the plant genera in
Table 1 , with the exception of Phyllostegia. However, none of these plant
genera have produced beach ecotypes on the present Hawaiian Islands, where
Phyllostegia is not known to occur below 300 m, and Pritctiardia is not known
below 3 0 0 m with the exception of a very small population on overhanging
basalt cliffs on the north coast of Molokai (Beccari & Rock, 1921). The Nihoa
population of Pritchardia occurs at 240 m, and the Tahiti population of
Phyllostegia (the only population known beyond the Hawaiian chain) is not
known to extend below 1200 m.
The Laysan populations of Himatione and Psittirostra represent the only
known occurrence of the endemic Hawaiian family Drepanidae on habitats
proximal to the beach. Their presence on Laysan lends very strong credence to
the concept of a relict, upland biota.
In their discussion of the land snail genus Tornatellides, Cooke & Kondo
(1960: 244) state: “The distribution of this genus differs completely from that
of any other genus of the family, in that it occupies most of the high islands
forming the northern, eastern, and southern peripheries of the islands of the
Pacific Ocean.”-“It is remarkable that not a single specimen of Tornatellides
has been taken on a single coral island in the south-central Pacific, although the
genus is abundant on the six low islands of Laysan, Lisianski, Pearl and Hermes,
Midway and Ocean (Kure) to the northwest of Hawaii and Johnston to the
southwest.” This evidence seems strong enough for the designation of the genus
as upland to montane, with a relictual status on the subsided islands of the
Leeward portion of the Hawaiian Chain. An additional land snail, Lamelliclea,
also occurs on Laysan. While it might be an upland relict, the widespread
214
S. 0. SCHLANGER AN11 G , W. GILLlrTT
occurrence of the genus on coral islands of the Pacific would seem to preclucie
such an interpretation.
The aspect of Laysan as a raised coral island 18,000 years ago may now be
observed in the high coral islands of Makatea and Henderson in the Tuamotu
Chain, each with a biota that includes land animals and upland to montane
plants. I t is significant that the biota of Makatea includes a contingent of land
snails (Cooke & Kondo, 1960) and the palm, Pritclzurtlh (Wilder, 1934), while
the biota of Henderson includes upland snails (Cooke & Kondo, 1960), a Rail,
Porzatiu (Olson, 1973), and a Sandalwood, Suritulzim (St John & Philipson,
1962), among othcr upland genera.
The interpretation of the nine lineages in Table 1 as upland to montane
relicts has been suggested by other biologists (Bitter, 1900; Carlquist, 1970),
and the geological and biological perspectives developed here are in close
agreement with those advanced by Zimmerman (1948). I t seems plausible that
the presence of these genera on Laysan derived from microevolutionary,
adaptive shifts concurrent with the invasions of the sea and subsidence of the
island. An alternative explanation that these portray recent ingression, via
long-distance dispersal, of lineages from upland habitats on distant high islands
appears to be less acceptable.
ACKNOWLEDGEMENTS
The support of this research by the U.S. National Science Foundation and
the Committee on Research, University of California, Riverside is gratefully
acknowledged. Special thanks are extended to E. D. Jackson, U.S. Geological
Survey, for his erudite and helpful review.
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