ATOLL RESEARCH BULLETIN

NO. 584

CHARACTERIZATION OF CORAL COMMUNITIES AT KINGMAN REEF IN

THE REMOTE CENTRAL PACIFIC OCEAN

BY

JEAN C. KENYON, JAMES E. MARAGOS, AND CASEY B. WILKINSON

ISSUED BY
NATIONAL MUSEUM OF NATURAL HISTORY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C., U.S.A.
DECEMBER 2010
Figure 1. Location of Kingman Reef in the central Pacific. Inset, center, shows IKONOS satellite imagery
of the atoll. Irregular white shapes overlaying the inset image are clouds.
 CHARACTERIZATION OF CORAL COMMUNITIES AT KINGMAN REEF IN
THE REMOTE CENTRAL PACIFIC OCEAN

BY

JEAN C. KENYON,1 JAMES E. MARAGOS,2 AND CASEY B. WILKINSON1

ABSTRACT

Kingman Reef, an atoll in the northern Line Islands of the central Pacific, is among
the last of the remote locations in the U.S. Pacific to be scientifically explored underwater,
and previously published surveys of its benthic reef communities have been limited to the
seaward- facing fore-reef habitat. Here, we apply multivariate statistical analyses to data
acquired in 2004 from several complementary survey methods that operate at different scales
of spatial and taxonomic resolution to characterize the coral communities in 5 habitats, 3 of
which are further stratified by geographic sector or depth zone. Coral cover is highest in the
small lagoon pool inside the southeast vertex of the perimeter reef, followed by the fore reef,
patch reefs, reef flat, and lagoon slope. Benthic communities are more highly variable on the
fore reef, but the coral cover and colony abundances are less dominated by a few key genera
than in other habitats. While the relative representation of coral taxa varies among habitats,
Porites, Fungia, Pocillopora, Sinularia, Turbinaria, Acropora, and Favia all contribute
> 10% of the total coral cover in at least one habitat. We provide a list of 188 anthozoan
and hydrozoan corals observed at Kingman Reef during survey activities since the year
2000. Coral diversity at Kingman is similar to that at Palmyra Atoll to the south but is high
compared to other adjacent atolls and reef islands to the south and west (Phoenix and Line
Islands) where data are available. Our spatially widespread surveys that generate independent
metrics of benthic cover and coral abundance collectively provide the most comprehensive
description of coral communities at Kingman Reef produced to date and also provide an
important record by which to monitor the response of this community to changing ocean
conditions.

___________________________________________________________________

1
Joint Institute for Marine and Atmospheric Research and NOAA Pacific Islands Fisheries Science
Center Coral Reef Ecosystem Division, 1125B Ala Moana Blvd., Honolulu, Hawai`i 96814, USA;
Email: [email protected]
2
Pacific Reefs National Wildlife Refuge Complex, U.S. Fish and Wildlife Service, 300 Ala Moana Blvd.,
Honolulu, Hawai`i 96850, USA

Manuscript received 10 November 2010; revised 1 December 2010.

2
INTRODUCTION

Kingman Reef (6o24′ N, 162o 24′W), located near the northern end of the Line
Island submarine ridge in the central Pacific (Fig. 1), is an atoll reef that encloses a deep
lagoon but lacks emergent land. First discovered in 1798, Kingman was named after an
American captain who next recorded its occurrence in 1853. Claimed over the following
80 years by businesses with interests in fishing and guano, Kingman Reef was placed
under the control of the U.S. Navy in 1934, which in turn transferred the atoll to the
U.S. Fish and Wildlife Service in 2001. Managed as a National Wildlife Refuge (NWR)
with boundaries extending to 12 nautical miles (nm) offshore, Kingman Reef NWR was
included within the Pacific Remote Islands Marine National Monument established by
President George W. Bush in January 2009, with monument boundaries extending 50 nm
from the mean low water line (U.S. Fish and Wildlife Service, 2010).
Roughly triangular in shape, the reef perimeter’s longest axis extends ~ 20 km in
an east-west direction with its acute eastern vertex facing into windward seas (Maragos et
al., 2008). The shallow (~ 3 m) eastern reef crest gradually deepens (~ 20 m) to the west,
and two small (~1–2 acre) emergent coral rubble ridges periodically shift contours atop
the eastern reef crest. About two dozen known pinnacles with patch reefs rise from the
lagoon floor in the shallower east and central area to within several meters of the surface
(Friedlander et al., 2010). Inside the southeastern vertex, a small (~ 0.14 km2 ), shallow
(maximum ~ 11 m) elliptical lagoon pool (Fig. 2b) hosts a dense assemblage of giant
clams and fungiid corals.
While known to support diverse and healthy marine communities, the absence of
habitable emergent land, coupled with its distance from human population centers, have
rendered Kingman Reef among the last of the remote locations in the U.S. Pacific to be
scientifically explored underwater. In 2000, reef assessment and monitoring activities
were initiated at Kingman Reef by the NOAA, Pacific Islands Fisheries Science Center’s
Coral Reef Ecosystem Division (CRED) in collaboration with scientists from the U.S.
Fish and Wildlife Service as part of a larger multidisciplinary effort to assess and monitor
coral reef ecosystems in the U.S. Pacific Islands (Brainard et al., 2008). Broad-scale
towed-diver surveys were initiated to provide a spatial assessment of the composition
and condition of shallow-water benthic habitats coupled with site-specific surveys to
assess species composition, abundance, percent cover, size distribution, and general
health of salient benthic organisms including corals. Beginning in 2002, biennial surveys
by CRED and its partners have continued to monitor major biotic and abiotic features
characterizing Kingman’s nearshore ecosystems so as to document temporal changes
and infer the processes underlying reef dynamics. Multidisciplinary studies of benthic
and fish assemblages in 2005 and 2007, led by the Scripps Institute of Oceanography, of
the fore - reef habitat at four atolls in the Line Islands experiencing varying degrees of
population density concluded that reef communities with relatively intact food webs such
as those at Kingman are the best available baselines for Pacific reefs (Sandin et al., 2008).
Several additional authoritative reports documenting the near-pristine nature of the coral
reef ecosystem at Kingman Reef have been provided by Brainard et al. (2005), Miller et
al. (2008), and Maragos et al. (2008).
 3
The main aims of the present study are to (1) describe the community structure
of the shallow-water (< 19 m) corals in 5 habitats at Kingman Reef, based on broad-
scale and site-specific quantitative surveys conducted in 2004, and (2) provide an
updated species lists of anthozoan and hydrozoan corals, based on surveys conducted
between 2000 and 2010. In describing community structure, we applied survey
methods that operate at different scales of spatial and taxonomic resolution to generate
independent metrics of community abundance. For each method and metric we applied
statistical analyses developed for use with multivariate ecological data to determine
spatial differences and their underlying taxonomic basis. The summarized results are
then integrated with the results of other studies that have examined coral communities
at Kingman Reef. Our study, which presents a detailed and spatially widespread
multivariate analysis of the coral communities at Kingman Reef, serves as an important
baseline from the early years of the 21st century, which can serve as an important and
useful standard for future generations of scientists, managers, and other stakeholders.

MATERIALS AND METHODS

Benthic Surveys

Towed-diver surveys were conducted in 2004 (2–4 April) according to the

methods of Kenyon et al. (2006a) in the fore-reef slope, lagoon slope, reef flat, and
eastern lagoon pool habitats (Figs. 2; 3a,b,c,e). Modifications to these methods included
the use of a digital still camera (Canon EOS-10D, EF 20-mm lens) in a customized
housing with strobes to photograph the benthos automatically at 15-sec intervals. Site-
specific belt-transect surveys to assess coral colony abundance and size class were
conducted by two divers (J. Kenyon and J. Maragos) from 2 to 4 April 2004, according
to the general methods described by Maragos et al. (2004) for the 2002 Rapid Ecological
Assessments. Two sites were surveyed on the fore-reef slope, 3 on the lagoon slope, 3
on hard-bottom patch reefs in the lagoon, and 1 in the eastern lagoon pool (Figs. 2; 3d,f).
Two, 25 x 2 m transect belts were surveyed at each site except for the eastern lagoon
pool, where only a single transect belt was surveyed as a result of the high density of
coral colonies. Each coral whose colony center fell within a 1- m-wide strip on each side
of the lines was classified to genus and its maximum diameter binned into one of seven
size classes (Mundy, 1996): 0–5 cm, 5–10 cm, 10–20 cm, 20–40 cm, 40–80 cm, 80–160
cm, or >160 cm. To determine percent cover of benthic components, 12 (35 cm x 50 cm)
photoquadrats were concurrently photographed with a Sony DSC P-9 digital still camera
with spatial reference to the same two 25-m transect lines: 3 photoquadrats at randomly
selected points along each transect and 3 at points 3 m perpendicular from each random
point in the direction of shallower water (i.e., 6 photoquadrats per transect; Preskitt et
al., 2004; Vroom et al., 2005). Locations of site-specific surveys were determined on
the basis of (1) maximizing spatial coverage within the allotted number of survey days;
(2) establishing depths that allowed three dives per day per diver; (3) working within
constraints imposed by other ship-supported operations; and (4) sea conditions.
4

Figure 2 . (a) Location of towed-diver and site-specific surveys at Kingman Reef in 2004. Tow track lines
are color coded according to the depth zone of the photographs sampled for image analysis. (b) Enlarged
view of the southeast corner better showing the eastern lagoon pool and reef flat survey areas.

Species lists of anthozoan corals, hydrozoan corals, and miscellaneous anthozoans

were compiled in the course of field surveys conducted between 2000 and 2010. Corals
were identified in situ to species (by the second author). Photographs were also taken in
situ for most species to verify taxonomic assignments and to resolve uncertainties using
published guidebooks. In rare circumstances, samples of some corals were also collected
to assist in identifications.
 5

A B

C D

E F

Figure 3 . Benthic communities in 5 habitats at Kingman Reef. (a) Fore-reef slope (b) Lagoon slope (c)
Reef flat (d) Lagoon patch reef (e, f) Eastern lagoon pool. Photos a, b, c, e acquired by towed-divers;
photos d, f acquired by roving divers.

Data Extraction and Analysis

Digital photographs from towed-diver surveys were sampled at 30-sec intervals

and quantitatively analyzed for benthic percent cover by a single analyst (C. Wilkinson),
using point-count software (Coral Point Count with Excel Extension; Kohler and
Gill, 2006) and using 50 stratified random points per frame. Benthic classification
categories included hard coral, soft coral, macroalgae, coralline red algae, non-coral
macroinvertebrates, sand, rubble, and pavement. Hard corals were subclassified
morphologically as branching, digitate, encrusting, massive, free-living (mushroom)
and plate-like. Laser-projected dots used to calibrate image size did not appear on
photographic imagery because of mechanical problems; consequently, scaling data
from imagery recorded throughout the Northwestern Hawaiian Islands with the same
6
methods (Kenyon et al., 2006b, 2007a, 2007b, 2008a, 2008b) were used to calculate
average benthic area in captured frames. Survey distances were calculated using a global
positioning system (GPS) and ArcView GIS 3.3. The depth of each captured frame was
determined by matching the time stamp of the image with that of temperature/pressure
data recorded by an SBE 39 recorder (Sea-Bird Electronics, Inc.) mounted on the habitat
towboard. For statistical analysis, each frame was categorized in one of 11 strata defined
a priori by the concatenation of 3 factors: habitat (fore reef, lagoon slope, reef flat, or
eastern lagoon pool), sector (northeast or south, where the southern margin of the eastern
lagoon pool was used to separate north from south) and depth zone (shallow [< 9.1 m] or
moderate [9.1–18.2 m]). Of 1581 frames, 66 (4.2%) were in depths > 18.2 m and were
eliminated from analysis as they were dispersed among 4 habitat sectors and may not
have sufficiently represented the spatially extensive deeper strata. The eastern lagoon
pool was not stratified by depth because of the small sample size (n = 12 frames), but
data were retained in statistical analyses because of the small area (~ 0.14 km2) of the
pool. Thus, frames from towed -diver surveys that spanned more than one habitat, sector,
or depth zone were parsed into separate strata using the time stamp that linked GPS
position and depth to recorded imagery.
Photoquadrats were analyzed for benthic percent cover by a single analyst
(J. Kenyon), using Coral Point Count with Excel Extensions (Kohler and Gill, 2006) and
using 50 stratified random points per digital image. Anthozoans were identified to genus,
while algae were categorized as Halimeda, other macroalgae, and coralline red algae.
Site locations and tracks of towed-diver surveys georeferenced with nondifferentially
corrected GPS units (Garmin® model 12) were mapped using ArcView GIS 3.3.
All statistical analyses were conducted using PRIMER-E®, version 6 (Clarke
and Warwick, 2001; Clarke and Gorley, 2006). For towed-diver surveys, benthic percent
cover data from each survey frame were treated as individual replicates within a stratum.
Two separate matrices were created. One matrix included percent cover data of all
benthic categories to examine differences in overall benthic composition among the
strata. A second matrix included only coral data, standardized as percent of total coral
cover, to examine differences in relative abundance of coral growth forms among the
strata. Square-root transformations were performed on the data matrices to lessen the
influence of prevalent benthic components and increase the weight of rare benthic
components, and Bray-Curtis resemblance matrices were created from transformed
data. To explore atoll-wide spatial distributional patterns, one-way analyses of similarity
(ANOSIM; maximum permutations = 9999) were conducted on each matrix to determine
if differences existed among strata. When ANOSIM detected global differences, R
values from pairwise tests were examined to determine where major differences existed.
SIMPER (similarity percentages routine) analyses using nontransformed data explored
the contribution of individual benthic components or coral groups to the dissimilarities.
To visually depict the relationships of strata to each other based on benthic
community structure, data were averaged by stratum, and a Bray-Curtis similarity matrix
(with square-root transformations) was generated to create a cluster diagram (group
average cluster mode) and nonmetric multidimensional scaling (nMDS) ordinations
(number of restarts = 50).
 7
Site-specific photoquadrat and coral belt-transect surveys were conducted
within the moderate -depth zone except for the eastern lagoon pool where surveys were
conducted at shallow depths (6.4–7.6 m). Percent cover data from each photoquadrat
image were treated as individual replicates within sites. Treatment of photoquadrat
percent cover data including all benthic categories was similar to that described for
towed-diver survey percent cover data. From a matrix including only coral data, ranked
cumulative abundance (k-dominance) curves were constructed from nonstandardized
data, and ANOSIM was conducted on a matrix of distances generated by DOMDIS
(dominance distance) between the dominance curves in the 4 habitats. To explore the
relative abundance of coral genera using the numbers of colonies rather than percent
cover as a metric, k-dominance curves were constructed from nonstandardized colony
count data, and ANOSIM was conducted on a separate matrix of distances generated by
DOMDIS between the k-dominance curves in the 4 habitats. R values in pairwise tests
were examined for both coral percent cover and colony count data to determine where
major differences existed. SIMPER analysis on nontransformed data explored average
similarities and dissimilarities among samples and the contribution of individual taxa to
the dissimilarities.
To examine differences in size-class structure among habitats of the two most
numerically abundant genera, Porites and Fungia, the number of colonies of each genus
was standardized and then cumulated for each transect at each site and a resemblance
matrix based on Euclidean distance was created for each genus. A one-way ANOSIM
was conducted for each genus to determine if the size-class distributions differed among
habitats.

RESULTS

Towed-diver Surveys

Survey Effort. The distance between frames sampled at 30-sec intervals from
benthic tow imagery depends on the tow speed; the average interframe distance ranged
from 17.0 m to 30.6 m (mean = 21.2 m, n = 21 tows). The average benthic area captured
in laser-scaled frames from the Northwestern Hawaiian Islands and applied to the
Kingman tows was 0.583 m2 (SE 0.01, n = 6398 frames). Towed divers surveyed 33.1
km of benthic habitat (Table 1, Fig. 2), from which 1515 frames were analyzed. Given
the 3:4 aspect ratio of the captured frames and extrapolating to the total number of
consecutive, nonoverlapping still frames that compose the benthic imagery, this benthic
analysis area (1515 frames x 0.583 m2/frame = 883 m2) samples a total survey area of
29,194 m2 (Table 1).
Survey effort, assessed by the area analyzed in sampled frames, was somewhat
greater on the fore-reef slope than the lagoon slope (434 m2 and 360 m2, respectively;
Table 1). Compared to shallow depths, survey effort at moderate depths was 2.5 times
greater on the fore-reef slope and 1.7 times greater on the lagoon slope. Of the 11 strata,
survey effort was greatest at moderate depths on the south fore-reef and northeast lagoon
slopes (186 m2 and 185 m2 analyzed in sampled frames, respectively).
8
Spatial Analyses of Benthic Cover and Composition. Percent cover estimates of
benthic components from analysis of towed-diver survey imagery exhibited 39% average
similarity among all fore-reef slope samples (n = 745), 48% average similarity among
all lagoon slope samples (n = 618), 52% average similar among all reef flat samples
(n = 140), and 54% average similarity among all eastern lagoon pool samples (n = 12)
(Table 1). Average dissimilarity among the 4 habitats was quite consistent, ranging from
62% (between the fore-reef slope and lagoon slope) to 68% (between the lagoon slope
and the reef flat). The fore-reef slope and eastern lagoon pool were dominated by coral
(39% and 62% of cover, respectively) (Table 1), while the lagoon slope was dominated
by turf-covered pavement (47% of cover) and the reef flat by coralline algae (49%
of cover). Average similarity within the 11 strata ranged from 34% to 62%, with the
lowest similarity found among shallow-depth south fore-reef samples and the highest
among shallow-depth northeast fore-reef samples (Table 1). On the fore-reef slope, the
highest coral cover (44%) was found in the shallow-depth south stratum, and the highest
coralline algal cover (49%) in the shallow-depth northeast stratum. On the lagoon slope,
the highest coral cover (25%) was found in the shallow-depth northeast stratum, and the
highest coralline algal cover (26%) in the shallow-depth south stratum.
One-way ANOSIM of towed-diver survey data revealed percent cover of benthic
components to be significantly different among the 11 strata (global R = 0.255,
p = 0.0001). Of the 55 pairwise tests, the greatest differences were found between the
eastern lagoon pool and the shallow northeast fore-reef strata (R = 0.65) and between the
southern lagoon slope and the northeast fore-reef strata at moderate depths ( R = 0.60)
(Fig. 4a). SIMPER tests on untransformed data revealed that a large suite of benthic
components contributed to the dissimilarities between the habitats, with turf-covered
pavement and coralline algae making the largest contributions.
Table 1. Survey effort, coralline algal and coral cover, and average similarity of benthic composition in sampled frames from
towed-diver surveys conducted at Kingman Reef, 2004. Shallow: < 9.1 m; Moderate: 9.1–18.2 m

Proportion of total coral cover

Mean %
Area Area
Distance a a
coralline Mean %
Depth surveyed surveyed # Frames analyzed algal cover total coral Average
2 2
Sector Zone (m) (m ) Analyzed (m ) (SE) cover (SE) Branching Digitate Encrusting Massive Mushroom Plate Octocorals Similarity

HABITAT: FORE-REEF SLOPE

ALL ALL 16930 14930 745 434 23.9 (0.8) 38.5 (0.9) 7.2 16.3 27.2 1.4 0.3 15.0 32.6 39.3
NE ALL 5669 4999 279 163 42.1 (1.1) 39.3 (1.2) 1.0 20.7 19.8 1.2 0.8 11.7 44.8 54.9
Shallow 63 37 49.1 (2.3) 31.0 (1.9) 1.0 21.7 31.9 0.8 1.2 8.6 34.7 61.7
Moderate 216 126 40.1 (1.2) 41.7 (1.4) 1.0 20.5 17.2 1.3 0.7 12.4 47.0 53.5
S ALL 11261 9931 466 272 13.0 (0.8) 38.0 (1.2) 11.1 13.5 31.8 1.5 0.1 17.1 25.0 38.7
Shallow 148 86 18.3 (1.6) 43.5 (2.3) 20.4 15.0 27.0 1.2 0.2 21.5 14.8 34.1
Moderate 318 185 10.6 (0.8) 35.4 (1.4) 5.7 12.7 34.5 1.6 0.0 14.6 30.9 42.6
HABITAT: LAGOON SLOPE
ALL ALL 13567 11964 618 360 15.9 (0.9) 20.1 (0.7) 2.1 5.5 56.8 4.7 8.7 10.5 11.7 47.8
NE ALL 7157 6312 365 213 10.5 (0.8) 22.1 (0.9) 1.2 2.9 56.7 3.3 7.9 12.6 15.3 52.3
Shallow 46 27 14.4 (0.3) 25.1 (2.7) 0.0 4.2 63.8 4.7 6.9 16.3 4.2 48.2
Moderate 319 186 9.9 (0.9) 21.7 (0.9) 1.4 2.7 55.5 3.1 8.1 12.0 17.1 53.4
S ALL 6410 5652 253 148 23.7 (1.6) 17.1 (1.1) 3.8 10.2 57.0 7.3 10.0 6.7 4.9 44.8
Shallow 183 107 26.1 (2.0) 17.4 (1.4) 4.1 13.0 52.3 9.3 10.2 7.1 3.9 42.9
Moderate 70 41 17.3 (2.2) 16.1 (1.7) 3.0 2.3 70.3 1.8 9.4 5.3 7.8 51.8
HABITAT: REEF FLAT
ALL ALL 2366 2087 140 82 48.6 (2.1) 30.3 (1.7) 1.4 17.6 42.0 11.0 15.7 6.4 6.0 52.0
NE Shallow 1497 1320 88 51 49.0 (2.9) 28.5 (2.2) 0.6 13.3 42.7 9.8 23.7 5.8 4.1 53.5
S Shallow 869 767 52 30 43.0 (3.0) 33.1 (2.7) 2.6 23.9 41.0 12.7 3.9 7.3 8.7 51.5
HABITAT: EASTERN LAGOON POOL
NE ALL 241 213 12 7 13.7 (4.5) 62.3 (5.5) 0.0 7.2 34.8 4.3 27.0 23.3 3.5 53.6

a
Area is based on average area of laser-scaled frames sampled at 30-sec intervals.
9
10
Spatial Analyses of Coral Cover and Composition. Mean total coral cover on
the fore-reef slope was 38.5% (SE 0.9, n = 745), 20.1% on the lagoon slope (SE 0.7, n =
618), 30.3% on the reef flat (SE 1.7, n = 140), and 62.3% in the eastern lagoon pool (SE
5.5, n = 12) (Table 1). Estimates of coral growth form relative abundance derived from
analysis of towed-diver survey imagery exhibited 39% average similarity among all fore-
reef slope samples, 47% average similarity among all lagoon slope samples, 40% average
similarity among all reef crest samples, and 56% average similarity among all eastern
lagoon pool samples. Average dissimilarity between samples from the 4 habitats ranged
from 57% to 68%, with the greatest average dissimilarities in coral growth form relative
abundance between the fore-reef slope and both the reef flat and the eastern lagoon pool
(data not shown).
On the fore-reef slope, octocorals dominated the northeast sector at both shallow
and moderate depths, while encrusting corals dominated the south sector in both depth
zones. Of the scleractinian corals on the northeast fore-reef slope, encrusting growth
forms were also the most abundant at shallow depths, but digitate forms were the most
abundant at moderate depths. The highest abundance of branching corals (primarily
Acropora) was found on the shallow south fore-reef slope. Massive and solitary
mushroom corals were rare on the fore-reef slope.
On the lagoon slope, encrusting corals dominated all exposures and depths,
accounting for more than half of the total coral cover. The next most abundant coral
growth form varied among each of the 4 strata of the lagoon slope (Table 1). Branching
corals were rare on the lagoon slope.
Encrusting corals were the most abundant growth form on the reef crest and in the
eastern lagoon pool. Solitary mushroom corals were the next most abundant growth form
in the eastern lagoon pool and on the northeast portion of the reef flat, but on the south
portion of the reef flat digitate corals were the second most abundant growth form.

Species Inventory of Corals and other Anthozoans

A total of 188 species of stony, soft, and hydrozoan corals and 11 other
anthozoans were recorded between 2000 and 2010 (Table 2). Within the class Anthozoa,
four taxonomic orders are represented. Within the stony coral order Scleractinia, 13
families containing 35 genera and 173 species are represented.

Site-specific Surveys: Photoquadrats

Spatial Analyses of Benthic Cover and Composition. Percent cover estimates

of benthic components from analysis of photoquadrat imagery exhibited 35% average
similarity among all fore-reef samples (n = 24) and 47% average similarity among all
lagoon samples (n = 84), but average dissimilarity between fore-reef and lagoon samples
was 65%. Average similarity within the 4 habitats ranged from 35% to 57%, with the
lowest similarity found among fore-reef samples and the highest among lagoon patch
reef samples. Scleractinian coral dominated the fore-reef slope and eastern lagoon pool
habitats, while pavement covered with turf algae dominated the inner lagoon slope and
lagoon patch reefs (Table 3).
 11

Figure 4. Nonmetric multidimensional scaling (nMDS) ordination depicting relationships of benthic

communities based on percent cover data. (a) Towed-diver surveys. Each symbol represents the average of
data derived from analysis of imagery captured at 30-sec intervals. Stratum key represents Habitat_Sector_
Depth, where FS = fore-reef slope, LS = lagoon slope, RF = reef flat, and PO = eastern lagoon pool for
habitat; S = south and NE = northeast for sector; S = shallow and M = moderate for depth.
(b) Photoquadrats. Each symbol represents a site where data from analysis of 12 photoquadrats were
averaged. LP = lagoon patch reefs.
12
Table 2. Scleractinian corals, octocorals, hydrozoan corals, and other Anthozoa at Kingman
Reef from 2000 to 2010 surveys and photos. Species assignments follow Veron et al.
(1977) and Veron (2000) except where there were conflicts with Hoeksema (1989) and
Wallace (1999) or contrary preferences of the authors (+). P = at Palmyra Atoll (Williams et
al. 2008)
SCLERACTINIAN CORALS A. expansa - P H. pilosa - P
Acanthastrea echinata A. listeri - P H. rigida - P
Acropora abrotanoides - P A. myriophthalma - P Isopora brueggemanni - P
Acropora aculeus - P A. suggesta Leptastrea bewickensis - P
A. acuminata - P Coscinaraea columna L. pruinosa - P
A. aspera - P Cycloseris cyclolites - P L. purpurea - P
A. austera Cycloseris sp. L. sp. [small dark round calices] - P
A. cerealis - P Cyphastrea decadia Leptastrea transversa - P
A.cf. clathrata - P C. ocellina Leptoria phrygia - P
A. cytherea - P Echinophyllia aspera - P Leptoseris mycetoseroides - P
A. digitifera - P E. echinata L. scabra
A. elseyi - P Favia favus - P Lobophyllia corymbosa - P
A. florida (polymorpha) - P F. laxa Lobophyllia hatai
A. gemmifera - P F. matthaii - P L. hemprichii (costata) - P
A. globiceps - P F. pallida - P Lobophyllia robusta
A. humilis - P F. rotumana - P Merulina ampliata - P
A. hyacinthus - P F. cf. rotumana Montastrea annuligera - P
A. latistella - P F. rotundata - P M. curta - P
A. longicyathus (syringodes) - P F. speciosa - P Montipora aequituberculata - P
A. microclados - P F. stelligera - P M. caliculata - P
A. monticulosa - P Favites abdita - P M. capitata - P
A. muricata (formosa) - P F. complanata M. danae - P
A. nana - P F. flexuosa - P M. efflorescens - P
A. nasuta - P F. halicora - P M. foliosa - P
A. paniculata - P F. pentagona - P M. flabellata - P
A. retusa F. russelli - P M. foveolata - P
A. robusta (conigera, nobilis)- P F. sp. M. grisea
A. rosaria - P Fungia concinna - P M. hoffmeisteri - P
A. samoensis - P F. corona M. cf. incrassata - P
A. sarmentosa F. danai - P M. informis - P
A. selago - P F. fungites - P M. millepora - P
A. sp.[undescribed Line&Phoenix]P F. granulosa - P M. monasteriata - P
A. sp. [undescribed] F. horrida (danai, klunzingeri)- P M. patula - P
A. sp. [slanted cones] F. paumotensis - P M. peltiformis - P
A. speciosa (rambleri) F. repanda - P M. sp. [encrusting]
A. spicifera - P F. scutaria - P M. tuberculosa - P
A. striata Gardineroseris planulata - P M. cf. turgescens
A. subulata - P Goniastrea edwardsi - P M. venosa - P
A. tenuis (africana) - P G. pectinata - P M. verrilli - P
A. valida - P Halomitra pileus - P Oxypora lacera
A. vaughani - P Herpolitha limax - P Pachyseris speciosa
A. verweyi - P H. weberi Pavona chiriquiensis - P
Alveopora verrilliana - P Hydnophora exesa - P P. clavus - P
Astreopora cucullata H. microconos - P P. duerdeni - P
 13

Table 2 (continued)

SCLERACTINIAN CORALS SCLERACTINIAN CORALS HYDROZOAN CORALS

P. explanulata - P P. vaughani - P Distichopora violacea - P
P. maldivensis - P P. sp. [smooth fingercoral] Millepora platyphylla - P
P. minuta - P P. sp. [large calices] Pennaria sp.
P. varians - P Psammocora haimeana - P Stylaster gracilis
P. venosa - P P. nierstraszi - P Unidentified hydrozoan coral
Platygyra contorta P. profundacella - P
P. daedalea (esperi) - P P. stellata - P Miscellaneous ANTHOZOA
P. lamellina - P P. superficialis Aplexidiscus sp. (Corallimorpharia)
P. pini - P Sandalolitha robusta - P Cryptodendrum adhaesivum (Actinaria)
P. ryukyuensis - P Scapophyllia cylindrica Discosoma sp. - P (Corallimorpharia)
P. sinenis- P Stylocoeniella armata Heteractis magnifica - P (Actinaria)
Plerogyra sinuosa Stylophora pistillata - P H. crispa (Actinaria) - P
Pocillopora brevicornis - P+ Turbinaria frondens - P H.malu (Actinaria)
P. capitata - P Turbinaria reniformis - P Phymanthus strandesi (Actinaria)
P. eydouxi - P T. stellulata - P Palythoa [large polyps] (Zoanthidea)
P. meandrina - P P. tuberculosa - P (Zoanthidea)
P. verrucosa - P OCTOCORALLIA CORALS Rhodactis howesii - P (Corallimorpharia)
P. zelli - P Cladiella pachyclados Stichodactyla mertensii (Actinaria)
Porites australiensis - P Dendronephthya mirabilis Zoanthus sp. (Zoanthidea)
P. evermanni - P Lobophytum sp.
P. lichen - P L. sp. B
P. lobata - P Pachyclavularia violacea - P
P. lutea - P Sarcophyton sp. - P
P. rus - P Sinularia sp. - P
P. pukoensis -stephensoni Stereonephthya unnicolor
P. solida - P Subergorgia suberosa
P superfusa - P+

Table 3. Mean percent cover of benthic components, derived from photoquadrat imagery at
Kingman Reef, 2004.
Lagoon
Category Fore reef inner slope patch reefs eastern pool
Scleractinian corals 43.4 22.0 31.6 66.0
Octocorals 8.0 0.6 0.6 0.7
Other Anthozoans 0.0 0.1 0.7 0.0
Halimeda 9.0 3.3 9.6 0.5
Other Macroalgae 1.6 0.4 0.4 0.2
Coralline Algae 11.0 4.0 14.3 2.5
Other Invertebrates 0.4 0.6 1.8 4.6
Pavement 20.6 44.6 39.2 22.9
Sand 4.6 15.2 1.8 2.4
Rubble 1.4 9.1 0.0 0.2

Average Similarity 35.2 49.7 56.7 51.3

14
One-way ANOSIM revealed percent cover of benthic components to be
significantly different among the 4 habitats (global R = 0.351, p = 0.0001). For all 6
pairwise comparisons, 0.278 ≤ R ≤ 0.410, indicating weak to moderate separation of
benthic components among habitats, with the strongest differences found between the
fore-reef slope and lagoon patch reefs, and between the eastern lagoon pool and the 2
other lagoon habitats (Fig. 4b). SIMPER tests on untransformed data confirmed that a
large suite of benthic components contributed to the dissimilarities between the habitats,
with Porites and turf-covered pavement making the strongest contributions.

Spatial Analyses of Coral Cover and Composition. Mean total coral cover in the
4 habitats ranged from 22.6% on the lagoon slope to 66.7% in the eastern lagoon pool
(Table 4). Estimates of coral relative abundance derived from analysis of photoquadrat
imagery exhibited 28% average similarity among all fore-reef samples and 51% average
similarity among all lagoon samples, but average dissimilarity between fore-reef and
lagoon samples was 74%.
Fifteen scleractinian and 4 alcyonacean genera (the octocorals Cladiella,
Lobophytum, Pachyclavularia, and Sinularia) were scored in photoquadrats (Table
4). In addition, the cnidarians Heteractis, Palythoa, and Stichodactyla were seen in
photoquadrat imagery but did not occur beneath randomly generated points. Average
similarity of coral relative abundance within the 4 habitats ranged from 28% to 57%
(Table 4), with the lowest similarity found among fore-reef samples and the highest
among lagoon slope samples. Cumulative ranked abundance (k-dominance) curves (Fig.
5a) demonstrated lower dominance (greater equitability) of coral genera in the fore-reef
habitat relative to the three lagoon habitats. ANOSIM revealed significant differences
among the k-dominance curves in the 4 habitats (global R = 0.507, p = 0.002), with the
greatest pairwise difference between the fore-reef and the lagoon slope habitats (R =
0.75) and the least between the lagoon slope and eastern lagoon pool (R = 0.1). SIMPER
revealed that Porites and Pocillopora together accounted for ~ 50% of the average
dissimilarity between the fore- reef and each of the 3 lagoon habitats, while Porites and
Fungia together accounted for ~ 63% of the average dissimilarity between each of the
lagoon habitats.
In all 4 habitats, Porites was the most highly ranked coral genus, with its highest
relative abundance on the lagoon slope and eastern lagoon pool (Table 4). Pocillopora,
Sinularia, and Acropora each accounted for a substantial proportion (> 10%) of the total
coral cover on the fore reef but were rare or absent in photoquadrat imagery in lagoon
habitats. Turbinaria was the second most abundant coral genus on the lagoon slope but
was absent from photoquadrat imagery in other habitats. Fungia was the second most
abundant coral genus on the lagoon patch reefs and the eastern lagoon pool. All other
genera accounted for < 10% of the total coral cover in all habitats.
 15
Table 4. Relative abundance of coral genera, expressed as percent of mean total coral
cover, derived from photoquadrat imagery at Kingman Reef, 2004.
Lagoon
Genus Fore reef inner slope patch reefs eastern pool
Acropora 13.0 0.0 0.0 0.0
Cladiella 0.0 0.0 0.4 0.0
Cyphastrea 0.2 0.2 1.9 0.0
Favia 4.3 0.2 2.6 2.3
Favites 0.2 0.0 0.9 0.5
Fungia 1.2 2.5 24.5 12.1
Gardineroseris 0.0 1.0 0.0 0.0
Goniastrea 0.0 0.0 0.6 0.0
Leptastrea 0.4 0.0 0.7 0.0
Lobophyllia 8.3 0.0 0.0 0.0
Lobophytum 0.0 2.5 0.7 0.5
Montastrea 0.0 0.0 0.4 0.0
Montipora 3.3 0.2 4.3 6.9
Pachyclavularia 1.4 0.0 1.1 0.3
Pavona 1.2 0.0 0.0 1.3
Platygyra 0.0 0.0 0.3 0.0
Pocillopora 18.3 3.5 0.0 0.3
Porites 34.2 75.3 61.7 75.6
Sinularia 14.2 0.0 0.0 0.2
Turbinaria 0.0 14.5 0.0 0.0

Mean (SE) coral cover,

% 51.4 (5.2) 22.6 (3.4) 32.2 (3.3) 66.7 (3.4)
Average Similarity 28.1 57.5 48.8 52.8
16

Figure 5. Cumulative ranked abundance curves (k-dominance curves) for corals in 4 habitats at Kingman
Reef. (a) Based on percent cover data from photoquadrats. (b) Based on colony counts in belt transects. FS
= fore-reef slope, LS = lagoon slope, LP = lagoon patch reefs, and PO = eastern lagoon pool.
 17
Table 5. Relative abundance of coral genera, expressed as percent of total number of
coral colonies, derived from belt-transect surveys at Kingman Reef, 2004.
Lagoon
Genus Fore reef inner slope patch reefs eastern pool
Acropora 4.4 0.1 0.9 0.2
Astreopora 0.0 1.9 0.7 0.0
Cladiella 5.6 0.2 0.9 0.1
Coscinaraea 0.1 0.0 0.0 0.0
Echinophyllia 0.2 0.8 0.5 0.0
Favia 11.4 4.7 9.3 2.4
Favites 2.0 0.9 2.3 1.0
Fungia 9.4 30.6 37.8 84.7
Gardineroseris 0.4 0.4 0.1 0.0
Goniastrea 0.0 0.0 0.2 0.0
Halomitra 0.1 0.0 0.0 0.0
Herpolitha 0.4 1.0 0.1 0.0
Hydnophora 1.3 0.5 0.1 0.1
Leptastrea 0.0 1.2 0.4 0.0
Leptoseris 0.1 0.0 0.1 0.1
Lobophyllia 0.7 0.0 0.0 0.0
Merulina 0.0 0.0 0.2 0.0
Millepora 0.0 0.0 0.1 0.0
Montastrea 0.3 0.9 1.3 0.2
Montipora 1.4 5.1 4.4 2.7
Pavona 2.7 0.5 1.5 0.3
Platygyra 1.6 0.2 0.3 0.1
Pocillopora 22.1 0.7 1.0 1.6
Porites 22.5 30.6 29.7 3.9
Psammocora 0.1 1.0 0.6 0.0
Sarcophyton 3.2 7.0 2.3 0.0
Sinularia/Lobophytum 7.1 9.3 4.5 2.6
Stylaster/Distichopora 0.2 0.0 0.0 0.0
Stylophora 0.6 0.0 0.0 0.0
Turbinaria 2.0 2.6 1.1 0.0

Total colonies
enumerated 941 1643 1930 1358
Average Similarity 59.7 51.9 49.5 —
18
Site-specific Surveys: Coral Belt Transects

Coral Colony Relative Abundance. A total of 5872 colonies belonging to at least

25 scleractinian, 3 alcyonace¬an (the octocorals Cladiella, Sarcophyton, and Sinularia/
Lobophytum), and 2 hydrozoan genera were recorded in belt-transects (Table 5). Average
similarity within the 3 habitats with multiple transects ranged from 50% to 60% (Table
5), with the lowest similarity found among lagoon patch reef samples and the highest
among fore-reef samples.
Cumulative ranked abundance (k-dominance) curves (Fig. 5b) demonstrated
lower dominance (greater equitability) of coral genera in the fore -reef habitat relative to
the 3 lagoon habitats. ANOSIM revealed significant differences among the k-dominance
curves in the 4 habitats (global R = 0.470, p = 0.001), with the greatest pairwise differ-
ence between the fore-reef and the eastern lagoon pool habitats (R = 0.98). Large R val-
ues, indicative of spatial differences in relative abundances of different coral taxa, were
also characteristic of the lagoon slope in comparison to both the eastern lagoon pool and
the fore -reef slope (R = 0.64 and 0.74, respectively). SIMPER revealed that Fungia and
Porites accounted for the greatest contributions to the dissimilarities between habitats.
At the 2 south fore-reef sites (Fig. 2), three genera — Porites, Pocillopora, and
Favia — accounted for more than half (56%) of all the colonies enumerated (Table
5, Fig. 5b). In the lagoon, Porites and Fungia accounted for more than 60% of all
the colonies enumerated in each of the slope and patch reef habitats, while Fungia
numerically dominated the eastern pool, accounting for 85% of enumerated colonies. All
other genera accounted for < 10% of the total number of colonies in all habitats.

Coral Size-class Distributions. Size-class distributions of Porites varied

significantly among the 4 habitats (global R = 0.332, p = 0.008) (Fig. 6). In pairwise tests,
the largest R value was associated with the comparison between the eastern lagoon pool
and the lagoon slope (R = 0.96). Inspection of size-class histograms for the 4 habitats
(Figs. 6a– d) reveals that less than 10% of the Porites colonies enumerated on the lagoon
slope had a maximum diameter > 40 cm, whereas more than 20% of Porites colonies
were in larger size classes (> 40 cm) in the other 3 habitats.
Inspection of size-class histograms for Fungia colonies suggests that the popula-
tion on the lagoon patch reefs was more strongly skewed towards smaller size classes
than populations in other habitats (Figs. 7a –d), but this trend was not statistically signifi-
cant (global R = 0.01, p = 0.40). All Fungia colonies had a maximum diameter < 40 cm.
 19

(a) Porites, Fore-reef Slope (b) Porites, Lagoon Slope

60
60
n = 212 n = 502
40 40

20 20

0 0
0‐5 5‐10 10‐20 20‐40 40‐80 80‐160 > 160 0‐5 5‐10 10‐20 20‐40 40‐80 80‐160 > 160

(c) Porites, Lagoon Patch Reefs (d) Porites, Eastern Lagoon Pool

60 60
n = 574 n = 53
40 40

20 20

0 0
0‐5 5‐10 10‐20 20‐40 40‐80 80‐160 > 160 0‐5 5‐10 10‐20 20‐40 40‐80 80‐160 > 160

Figure 6(a–d). Size -class (cm) distributions of Porites in 4 habitats at Kingman Reef. The x-axis is
maximum diameter (cm); the y-axis is percent of Porites colonies enumerated in each habitat.

(a) Fungia, Fore-reef Slope (b) Fungia, Lagoon Slope

60 60
n = 88 n = 503
40 40

20 20

0 0
0‐5 5‐10 10‐20 20‐40 40‐80 80‐160 > 160 0‐5 5‐10 10‐20 20‐40 40‐80 80‐160 > 160

(c) Fungia, Lagoon Patch Reefs (d) Fungia, Lagoon Eastern Pool

60 60
n = 729 n = 1150
40 40

20 20

0 0
0‐5 5‐10 10‐20 20‐40 40‐80 80‐160 > 160 0‐5 5‐10 10‐20 20‐40 40‐80 80‐160 > 160

Figure 7(a–d). Size -class (cm) distributions of Fungia in 4 habitats at Kingman Reef. The x-axis is
maximum diameter (cm); the y-axis is percent of Fungia colonies enumerated in each habitat.
20

DISCUSSION

Previous quantitative descriptions of benthic community structure and coral

cover at Kingman Reef are limited to the fore-reef slope habitat, based on surveys
conducted in 2005 at 10 sites (Sandin et al., 2008). Here, we apply multivariate statistical
analyses to data acquired in 2004 from several complementary survey methods that
operate at different scales of spatial and taxonomic resolution to characterize the
benthic communities at Kingman Reef in relationship to strata defined by habitat,
geographic sector, and depth zone. Analysis of imagery recorded during broad-scale
surveys conducted by towed divers provide a spatially extensive assessment of benthic
communities in the fore-reef slope, lagoon slope, reef flat, and eastern lagoon pool
habitats at a relatively coarse level of taxonomic resolution, while site-specific surveys
using photoquadrats and belt transects in the fore-reef slope, lagoon slope, lagoon patch
reef, and eastern lagoon pool habitats enable finer resolution of corals to the genus or
species level. In the following discussion, the diverse metrics generated by these three
methodologies are synthesized to provide a baseline description of coral communities at
Kingman Reef.
The separation of benthic assemblages among the 4 habitats surveyed by each
methodology is graphically demonstrated by nonmetric multidimensional scaling (nMDS)
plots, based on data extracted from towed-diver survey and photoquadrat imagery (Fig.
4). For both methodologies, a large suite of benthic components contributed to the
dissimilarities among habitats, rather than being driven by a few components. Benthic
communities in the fore-reef habitat were distinguished from those in other habitats by
the lowest among-sample similarities (i.e., highest variabilities) generated from both
broad-scale and photoquadrat survey data. Site-specific photoquadrat and belt-transect
surveys demonstrated that coral assemblages on the fore-reef slope had lower dominance
(i.e., greater equitability) of coral genera than assemblages in the 3 lagoon habitats
(slope, patch reefs, eastern pool). Thus, while the fore-reef samples showed the greatest
dissimilarity to each other, the fore-reef coral assemblage was less dominated by a few
principal genera than was the case in other habitats. Mean coral cover in the 4 fore-reef
strata surveyed by towed divers was 38.5%.
Of the 11 strata surveyed by towed divers, the lowest similarity (34.1%) was
found among samples representing the shallow south fore reef. The shallow south fore
reef was also distinguished from other fore-reef strata by hosting the highest mean coral
cover (43.5%). The highest abundance of branching corals (primarily Acropora, data not
shown) was on the shallow south fore reef. Sandin et al. (2008) also noted the abundance
of branching corals on the fore reef, especially Acropora. Nonetheless, encrusting corals
were the most abundant growth form on the south fore reef at both shallow and moderate
depths.
Sandin et al. (2008) report a mean scleractinian (stony) coral cover ~ 42% from
image analysis of photoquadrats at 10 fore-reef sites between depths of 10 and 12 m in
the south and northeast sectors at Kingman Reef. In contrast, mean scleractinian coral
cover from towed-diver surveys between depths of 10 and 12 m on the fore reef (n = 175
frames) was 26.1% (SE 1.4) (data not shown). Estimates of coral cover derived from site-
 21

specific surveys generally exceed those derived from towed-diver surveys (Kenyon et al.,
2006b, 2007a,b, 2008 a,b), because site-specific surveys target hard-bottom communities,
whereas broad-scale surveys include more variable substrate. Our estimate of coralline
algal cover from towed-diver surveys between depths of 10 and 12 m on the fore reef
(18.8%, SE 1.5) also differs from that reported by Sandin et al. (2008), ~ 29%. In the
present study, coral cover reported from photoquadrats at 2 sites on the south fore reef
at moderate depths was 51.4%, whereas coral cover from broad-scale surveys in this
stratum was 35.4%. While the capacity to distinguish among coral taxa in photoquadrats
is superior to that in towed-diver survey imagery, the much larger area analyzed from
towed-diver surveys suggests that this method generates the more accurate estimate of
total coral cover over broad spatial scales. In the present study, nearly twice as many
coral genera were recorded in fore-reef belt-transect surveys (25) than in photoquadrats
2 2
(13), which is not unexpected given the comparative areas surveyed (200 m vs. 4.2 m ).
The two northeast fore-reef strata were most unlike other strata in terms of overall
benthic composition (Fig. 4a). Octocorals dominated the northeast fore-reef slope in both
depth zones, accounting for more than a third of the total coral cover (Table 1).
Estimates of coral cover from broad-scale surveys of the lagoon slope ranged
from 16.1% in the south sector at moderate depths to 25.1% in the northeast sector
at shallow depths, with a habitat-wide average of 20.1%. In all lagoon slope strata,
encrusting coral was the most dominant growth form (Table 1). Estimates of coral cover
from photoquadrat imagery in the lagoon slope habitat (22.6%, Table 3) are in good
agreement with those from broad-scale surveys. Porites made the greatest contribution
to total coral percent cover (75.3%; Table 4, Fig. 5a) and, with Fungia, co-dominated
the coral assemblage in regards to the number of colonies (Table 5, Fig. 5b). Despite the
abundance of Porites in the lagoon slope habitat, the population was distinguished from
congeneric populations in other habitats by a smaller proportion of colonies in large (>
40 cm maximum diameter) size classes (Fig. 6b). Turbinaria was the next most abundant
coral genus in terms of percent cover (14.5%) but accounted for less than 3% of the total
number of colonies enumerated. Average similarity values among lagoon slope samples
acquired by different methodologies were quite consistent, ranging from 48% (Table 1)
to 52% (Table 5), with the exception of higher average similarity (i.e., less variability) of
coral relative abundance samples from photoquadrats (58%; Table 4). A general statement
by Maragos et al. (2008) that “corals are more abundant and varied on seaward [fore-
reef] slopes vis-à-vis lagoon slopes” is supported by percent cover data (Tables 1, 3) and
k-dominance curves (Fig. 5) in the present study.
To date, the shallow reef flat has only been surveyed with the use of towed-diver
surveys. This method is well suited to assess habitat that may be difficult for roving
divers to survey due to strong surge or currents. Coral cover was relatively high in this
habitat, averaging 30.3% over the 2.4 km surveyed (Table 1). The most dominant coral
growth form was encrusting, with mushroom corals abundant on the northeast reef flat
adjoining the eastern lagoon pool.
Benthic components in samples from lagoon patch reefs showed the highest
among-sample similarity (56.7%) of the 4 habitats surveyed with photoquadrats.
Mean coral cover was 32.2%, and mean coralline algal cover was greater in this
22
habitat than the other habitats surveyed by roving divers (14.3%). Porites made the
greatest contribution to coral percent cover (61.7%, Table 4) and accounted for ~30%
of the colonies enumerated in this habitat, with moderately sized colonies (20–40 cm
maximum diameter) as the most abundant size class. Fungia was the most abundant
genus in terms of the number of colonies (37.8% of total) and also accounted for nearly
a quarter of the total coral percent cover, although few Fungia colonies exceeded 20
cm maximum diameter. Coral dominance patterns were intermediate between those
demonstrated in the fore-reef slope and eastern lagoon pool habitats.
Of the 5 habitats surveyed, mean coral cover was highest in the eastern lagoon
pool, with good agreement between values generated from broad-scale surveys (62.3%,
Table 1) and from photoquadrats (66.7%, Table 3). Encrusting corals were the most
dominant growth form in broad-scale survey imagery, followed by mushroom corals
(34.8% and 27.0% of coral cover, respectively). Porites only accounted for 3.9% of
the total number of colonies enumerated (Table 5), but made the greatest contribution
to percent cover (75.6%; Table 4). Conversely, Fungia accounted for 84.7% of the
colonies enumerated at the site surveyed by roving divers (Fig. 2b, Table 5), yet only
contributed 12.7% of the total coral cover (Table 4). Maragos et al. (2008) also note the
large number of Fungia colonies in the eastern lagoon pool. Colony abundance, derived
from belt transects, and percent cover, derived from photoquadrats, are two independent
metrics of coral relative abundance. A taxon with a small number of large colonies may
contribute substantially to total coral cover but only account for a small proportion of
the total number of colonies in a coral assemblage. A quarter of all the Porites colonies
enumerated in the eastern lagoon pool were very large (80–160 cm maximum diameter;
Fig. 6d) which helps to explain the disparity between the number of colonies and percent
cover. Similarly, the lack of correspondence in Fungia between relative abundance as
indexed by percent cover and by proportion of colonies is driven by the comparatively
small size of Fungia colonies (Fig. 7d). Both percent cover and colony abundance are
important metrics. Percent cover, along with growth morphology, shapes the physical
structure of the benthic environment experienced by organisms such as fish and other
invertebrates (Jones et al., 2004; Idjadi and Edmunds, 2006), while colony abundance
influences density-dependent processes including disease transmission by infectious
vectors, the probability of fertilization during sexual reproduction, and habitation for
commensal organisms that occupy different coral taxa.
Of 199 cnidarian taxa reported from Kingman Reef in this study, 147 (73.9%)
have also been reported
o
from oPalmyra Atoll (Williams et al., 2008; J. Maragos, unpubl.
data), centered at 5 52′N, 162 06′W to the south. Conversely, of 198 cnidarian taxa
reported from Palmyra by Williams et al. (2008) and Maragos (unpubl. data), 141
(71.2%) are reported in this study. Four scleractinian genera observed at Kingman Reef
(Acanthastrea, Coscinaraea, Plerogyra, Stylocoeniella) have not been reported from
Palmyra, and 6 cnidarian genera including 5 scleractinian genera observed at Palmyra
(Balanophyllia, Ctenactis, Plesiastrea, Symphyllia, and Tubastraea) have not been
reported from Kingman. Kingman and Palmyra lie predominantly in the North Equatorial
Countercurrent (NECC), a warm, eastward flowing surface countercurrent that seasonally
varies in position and strength. During the boreal spring (February–April), the NECC is
 23
-1 o
relatively weak (0.2 m s ) and-1
restricted to 4–6 N, but from May–January,
o
NECC surface
flow increases (0.4–0.6 m s ) and widens to lie between 5–10 N (Maragos et al., 2008).
This current flow from the west may carry the larvae of corals and other reef life from
the biodiversity-rich western Pacific to central Pacific reefs influenced by the NECC. Of
10 reef systems surveyed in the Phoenix (Baker, Howland, Kanton, McKean) and Line
Islands (Kingman, Palmyra, Teraini, Tabuaeran, Kiritimati, Jarvis), the highest numbers
of species and genera of stony corals have been reported from Kingman and Palmyra
(Maragos et al., 2008). In turn, Kingman may serve as a source of larvae to other, distant
reef systems. Genotype analysis of 7 microsatellite loci from a rare colony of Acropora
cytherea off Kauai in the main Hawaiian Islands (Kenyon et al., 2007c) corresponded
more closely to samples of A. cytherea from Kingman Reef, ~1750 km distant, than to
conspecific samples from French Frigate Shoals in the Northwestern Hawaiian Islands
or Johnston Atoll, ~ 1200 km and 700 km distant, respectively (G. Concepcion, unpubl.
data).
Larval dispersal affects the distribution and abundance of marine organisms,
enabling the replenishment of populations, colonization of new habitats, and expansion
of geographic ranges. Significant controversy exists about the scale of coral dispersal,
in particular whether populations are mostly self-seeded (i.e., most successful larvae
recruit to their natal reef) or are maintained principally by recruits arriving from nearby
or even distant reefs (Strathmann et al., 2002). For corals, knowledge about the survival
dynamics of larvae, even under laboratory conditions, is sparse. Prior to studies by
Graham et al. (2008), estimates for maximum survival times for broadcast-spawning
scleractinian corals ranged from 23 to 130 days. In laboratory cultures, Graham et al.
(2008) quantified the survival of larvae from 5 broadcast-spawning coral species with
lecithotrophic larvae (i.e., they cannot supplement their maternal energy reserves but
only acquire zooxanthellae after settlement and metamorphosis) and tested for changes
in mortality rates as larvae aged. Maximum species-specific lifespans ranged from 195
days to 244 days. Three periods were identified: high initial rates of mortality; followed
by a low, approximately constant rate of mortality; and finally, progressively increasing
mortality after approximately 100 days. Because coral larvae are relatively poor
swimmers, their dispersal distances will largely depend on the duration of the pelagic
phase and the speed and direction of water currents transporting the larvae (Scheltema,
1986). The observed lifetimes suggested that the potential for long-term dispersal of coral
larvae may be substantially greater than previously thought, and may partially explain the
large geographic ranges of many species (Hughes et al., 2002). Detection of increasing
mortality rates late in life suggests that energy reserves do not reach critically low levels
until approximately 100 days after spawning. Conversely, increased mortality rates early
in life decrease the likelihood that larvae transported away from their natal reef will
survive to reach nearby reefs, and thus decrease connectivity at regional scales.
The capacity of larvae to disperse between distant reefs and Kingman Reef
appears favorable, and local populations of some taxa are known to be reproductively
capable. Kenyon (2008c) examined developing eggs in 7 species of Acropora from
Kingman Reef in 2002 and 2004 and, by comparing the sizes of developing eggs to the
size of mature eggs reported in scientific literature, inferred several episodes of spawning,
24
each involving multiple species, taking place over 2 or 3 months beginning in early
spring. Recruitment and growth of coral larvae also appears robust at Kingman Reef.
Over a 6-year study in which cohorts of settlement plates were deployed for periods of 2
years at Kingman, Palmyra, Jarvis, and Baker in the Line-2
and
-1
Phoenix Islands, the highest
average annual recruitment rate (8.1 ± 2.8 recruits m year ) was observed at Kingman
Reef (Kenyon, 2009). The taxonomic composition of the recruits varied among cohorts,
although the settlement plates were deployed at the same location on the shallow (7.9 m
depth) southeast lagoon slope at the same time of year for equal periods of time, with the
first cohort dominated by Acropora recruits (80% of total) and the final cohort dominated
by Porites (90% of total). Rapidly growing acroporids from the first cohort averaged 24.6
cm geometric mean diameter while more slowly growing poritids from the final cohort
averaged 7.6 cm geometric mean diameter (J. Kenyon, unpubl. data). Recruitment to the
middle cohort deployed from 2004 to 2006 was among the lowest rate documented in
all the remote Pacific locations during the 6-year study, and the reasons for this failure
in recruitment are unclear. At Palmyra Atoll, densities of small (< 5 mm) juvenile corals
examined in situ were 9 times higher on the fore reef than on the back reef (Roth and
Knowlton, 2009), suggesting recruitment rates may be higher at Kingman Reef on the
fore-reef slope than the lagoon slope habitat. The temporal and taxonomic variability of
recruitment is one of several biotic processes including growth, competition, disease, and
predation that influence the structure of coral communities.
Kingman has been referred to as a pristine reef (Friedlander et al., 2010) because
of unaltered fish assemblages and the relative absence of anthropogenic impacts, though
other authors contend “there are no pristine reefs left” (Hughes et al., 2003). Some
threats such as climate change are very widespread and will likely challenge even the
capacity of reefs that are distant from population centers to remain ecologically intact
(Hoegh-Guldberg and Bruno, 2010). As the number of reports of degrading coral reefs
increases in response to local and global stressors (e.g., Knowlton, 2001; Hughes et al.,
2003; Pandolfi, 2005; Hoegh-Guldberg et al., 2007), it becomes increasingly urgent
to establish detailed descriptive baselines by which the direction and pace of future
trajectories can be determined. Our application of multivariate statistical analyses to
survey data collected at different scales of spatial and taxonomic resolution has discerned
spatial variation that provides a baseline for future surveys that may range in their
capacity for spatial coverage and the metrics generated.

ACKNOWLEDGMENTS

This work is part of an interdisciplinary effort by the NOAA Pacific Islands

Fisheries Science Center Coral Reef Ecosystem Division (CRED) to assess and monitor
coral-reef ecosystems in the U.S. Pacific in collaboration with marine scientists at
the U.S. Fish and Wildlife Service. We thank the officers and crew of the NOAA
Ships Oscar Elton Sette and Hi`ialakai for logistic support and field assistance.
Molly Timmers and Kyle Hogrefe assisted in the collection of towed-diver survey
data; Kimberly Page and Linda Preskitt collected photoquadrat data. Permission to
work at Kingman Reef was granted by the Pacific Remote Islands Wildlife Refuge
 25
Complex, U.S. Fish and Wildlife Service, Department of the Interior. Funding from
NOAA’s Coral Reef Conservation Program and from the U.S. Fish and Wildlife Service
supported this work.

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