A Journal of Conservation Biogeography Diversity and Distributions, (Diversity Distrib.

) (2015) 21, 465–476

BIODIVERSITY The emergent geography of biophysical

RESEARCH
dispersal barriers across the Indo-West
Pacific
Eric A. Treml1*, Jason Roberts2, Patrick N. Halpin2, Hugh P. Possingham3
and Cynthia Riginos3

1
School of BioSciences, University of ABSTRACT
Melbourne, Melbourne, Vic., 3010, Australia,
2 Aim To discover and evaluate potential dispersal barriers across the Indo-West
Marine Geospatial Ecology Laboratory,
A324 LSRC Building, Nicholas School of the
Pacific Ocean and to develop spatially explicit hypotheses regarding the loca-
Environment, Duke University, Durham, tion of barriers and their capacity to filter taxa. Additionally, to compare model
NC, USA, 3School of Biological Sciences, The predictions with previously described barriers and build a more thorough
University of Queensland, St. Lucia, Qld, understanding of the region’s biogeographic patterns.
4072, Australia Location The reefs of the Indo-West Pacific Ocean, from 100 to 170°E and
from 30°N to 30°S.

Methods A biophysical larval dispersal model was used to quantify over

800,000 dispersal kernels by simulating 99 model taxa. Spatial analysis was used
to map emergent patterns of potential multispecies population connectivity and
predict geographic locations and strength of concordant dispersal barriers
across the seascape.

Results Although individual taxa varied in their predicted fine-scale population

connectivity, there was consistency in the location of dispersal barriers. Across
all trait combinations, reproductive output (fecundity and density of reproduc-
Diversity and Distributions

tive adults), reproductive strategy (spawning phenology) and the length of the
larval dispersal phase determined the strength of barriers across the seascape.
More than 80% of the previously described marine boundaries in the region
were predicted to restrict the dispersal of at least 10% of the simulated taxa.
The location and strength of many additional multitaxa barriers were described
at scales <200 km.

Main conclusions The geographic structure of many previously described mul-

*Correspondence: Eric A. Treml, School of
BioSciences, University of Melbourne,
Melbourne, Vic. 3010, Australia.
E-mail: etreml@unimelb.edu.au
tispecies barriers across the Indo-West Pacific were predicted with the biophysi-
cal dispersal model and spatial analysis. The permeability of individual
dispersal barriers was primarily determined by reproductive output, spawning
phenology and the length of the pelagic larval stage. Across all dispersal barriers
analysed, Euclidean distance explained up to 49% of the variation in barrier
strength.
Keywords
Barriers, biophysical modelling, connectivity, coral reef, dispersal, network
analysis.

systems, the identification of barriers, and the processes that

INTRODUCTION
Although barriers to dispersal are known to contribute to
the distinctiveness of communities, species and populations,
the importance of the biophysical processes associated with
permeable barriers is not well understood (Mayr, 1963;
MacArthur & Wilson, 1967; Bowen et al., 2013). In marine
create or maintain them, is confounded by the dynamic
nature of the physical environment and the diversity of bio-
logical traits evolved to thrive in these environments. Many
marine organisms disperse only as larvae, thus population
connectivity is largely determined by biological traits and
the physical environment (Cowen & Sponaugle, 2009) and

DOI: 10.1111/ddi.12307
ª 2015 John Wiley & Sons Ltd http://wileyonlinelibrary.com/journal/ddi 465
E. A. Treml et al.
complex interactions between biology and barriers (Ayre
et al., 2009). Although determinants of broad-scale marine
population connectivity include egg type (Bradbury et al.,
2008; Riginos et al., 2011), larval mortality (Cowen et al.,
2000), pelagic larval duration or PLD (Shanks, 2009), larval
output (Steneck et al., 2009; Treml et al., 2012), geography
(Bradbury et al., 2008; Riginos et al., 2011) and the near-
shore environment (Largier, 2003), relatively little is known
regarding the broad-scale biogeographic patterns resulting
from this connectivity. In addition, marine organismal
diversity and geographic variability in the seascape lead to
broad differences in population connectivity across taxa
(Kinlan & Gaines, 2003; Gaither & Rocha, 2013), making a
synthesis difficult (Werner et al., 2007; Cowen & Sponaugle,
2009).
Marine population connectivity should influence patterns
in phylogeography and population genetics (Palumbi, 1994),
species diversity (Hobbs et al., 2010), isolation and ende-
mism (Meyer et al., 2005) and metapopulation persistence
(Hastings & Botsford, 2006). The localized process of a sin-
gle dispersal event has both short-term demographic conse-
quences and, when aggregated across generations, long-term
implications for genetic differentiation (Palumbi, 1994; Rigi-
nos et al., 2011). In general, as the dispersal of individuals
between populations increases, gene flow increases, and
genetic differentiation between populations is diminished
(Slatkin, 1987). In contrast, as population connectivity
decreases, genetic differentiation is expected to increase and
populations spanning dispersal barriers become increasingly
isolated (Mayr, 1963). With increased isolation, there
should be an increase in endemic species (MacArthur &
Wilson, 1967; Meyer et al., 2005), lower total species diver-
sity (Sandin et al., 2008) and lower population resilience
(Hastings & Botsford, 2006). Because population connectiv-
ity is expected to have such profound effects on population
resilience and diversity, it remains a fundamental concern
in marine conservation efforts (Palumbi, 2004; Jones et al.,
2007).
Dispersal barriers may develop through any combination
of: (1) biological constraints such as larval type, mortality
and competency characteristics (e.g. Cowen & Sponaugle,
2009); (2) historical or geologic barriers such as land bridges
(Hoeksema, 2007); (3) ocean currents or eddies (e.g. Barber
et al., 2006); (4) environmental limitations (Marshall et al.,
2010); or (5) distance between habitat patches (e.g. Lessios &
Robertson, 2006). Adding to this biophysical complexity is
the variable time-scales at which these processes may be act-
ing (Riginos & Liggins, 2013). Strong persistent barriers such
as the Isthmus of Panama (Knowlton & Weigt, 1998) or the
Eastern Pacific Barrier (Lessios & Robertson, 2006) may have
led to divergence over millennia. However, other boundaries
are sites of previous vicariance, such as the Sunda Shelf break
(Randall, 1998) and the Torres Strait (Mirams et al., 2011),
now regions with minimal contemporary dispersal limita-
tions. Examples of inferred contemporary dispersal barriers
include the Mona Passage for a coral (Baums et al., 2005)
466
and the Halmehera Eddy for stomatopods (Barber et al.,
2006). Because there are few absolute barriers to dispersal in
the present-day ocean, many so-called barriers may be semi-
permeable with taxon-specific responses affected by biologi-
cal traits (Ayre et al., 2009). As a result, dispersal studies
have shifted towards understanding these biological traits,
and the ecological and evolutionary processes that lead to
barriers in the ocean (Mora et al., 2003; Bradbury et al.,
2008; Riginos et al., 2011; Treml et al., 2012).
One region of intensive connectivity research is the biodi-
versity hotspot of the Coral Triangle within the Indo-West
Pacific (IWP) Ocean; the seascape encompassing the Philip-
pines, Malaysia, Indonesia, Timor-Leste, Papua New Guinea
and the Solomon Islands (Veron et al., 2009). This region is
of global importance for biodiversity (Roberts et al., 2002),
economic benefits and human livelihood (Burke et al.,
2012). Although a more comprehensive picture is emerging
in the IWP with respect to marine phylogeographic and pop-
ulation genetic patterns (Crandall et al., 2008, 2014; Timm &
Kochzius, 2008; Carpenter et al., 2011) and the centre of bio-
diversity (Hoeksema, 2007; Gaither & Rocha, 2013), the spa-
tial structure and processes affecting dispersal across species
is poorly understood.
Few methods exist for developing a geographically and
biologically comprehensive synthesis of marine dispersal bar-
riers. Exceptions include combined data analysis (Bradbury
et al., 2008; Riginos et al., 2011), comparative phylogeo-
graphy (Lessios & Robertson, 2006; Hickerson & Meyer,
2008; Ayre et al., 2009) and biophysical dispersal modelling
(Cowen et al., 2000; Werner et al., 2007; Kool et al., 2011).
Recently, biophysical modelling studies of larval dispersal
have successfully quantified potential ecological connectivity
(Cowen et al., 2006; Werner et al., 2007) and accurately pre-
dicted genetic structuring (e.g. Galindo et al., 2006; White
et al., 2010; Davies et al., 2014), but have not resolved the
geographic patterns and characteristics of concordant
dispersal barriers across taxa.
In this study, we predict the geographic location and
strength of multitaxon dispersal barriers across the Indo-
West Pacific Ocean. We use empirically based life history
parameters and a regional ocean model (Treml et al., 2008,
2012) to simulate dispersal and identify barriers. In previous
studies, we used this model to explore ecological connectivity
patterns among political units (Treml & Halpin, 2012) and
completed a comprehensive sensitivity analysis (Treml et al.,
2012). Here, we expand on this model and consider a much
greater range of biological traits to predict the locations and
strengths of multitaxon barriers across the IWP. Specifically,
we ask three questions: (1) What biological attributes are
important in influencing the location and strength of dis-
persal barriers? (2) What is the geographic configuration of
multitaxon dispersal barriers? (3) Is there geographic concor-
dance with previously described and empirically based barri-
ers? Our results provide detailed predictions that can be
compared to future empirical studies of biodiversity
patterns.
Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd

METHODS
A dispersal model (Treml et al., 2012) was modified to simu-
late population connectivity across a range of biologically
realistic traits for the entire Indo-West Pacific (IWP),
extending from 100 to 170°E and 30°N to 30°S (Fig. 1). The
model seascape was parameterized using high-resolution
coral reef habitat data (Spalding et al., 2001) and surface
currents (~top 10 m) from the 1/12° eddy-resolving Regional
Ocean Modeling System – ROMS (Wang & Chao, 2004;
Wang et al., 2005). All spatial data were interpolated to the
10 by 10 km model grid, consistent with the resolution of
the hydrodynamic data. To maximize computational effi-
ciency and capture intra- and interannual variability, we sim-
ulated dispersal for each trait combination across three
specific years: 1997, a strong El Ni~
no year; 1999, a strong La
Ni~na year; and 2001, a neutral year. Simulations were aggre-
gated to quantify potential connectivity within a ‘typical’
5-year span, assuming an El Ni~ no and La Ni~ na frequency of
two events per decade (Quinn et al., 1993). The model’s res-
olution and extent allow the downstream dispersal kernel to
be quantified, but probably underestimate local-scale reten-
tion below the 10 km model resolution.
With the intent to explore the full range of dispersal capa-
bilities across marine organisms, we created 99 hypothetical
taxa (Appendix S1 in Supporting Information). Each taxon
was represented by a unique combination of life history
characteristics known to influence connectivity (reviewed by
North et al., 2009), namely: spawning periodicity (seasonal
(A)
Philippines
15o
(B) (C)
Guam
(D)
Palau
(E)
Halmahera
Cenderwasih Bay
(H)
0o (I)
(F)
(G)
Solomon
PNG
Indonesia
–15o
(L)
Australia
Ningaloo
105oE 120o 135o 150o
Figure 1 Study region. The Indo-West Pacific Ocean with
geographic features and previously described barriers depicted
(see Table 2 and text for description of barriers). Model sample
sites are shown as black points and the coastal shelf to the
120 m isobath is shaded. All maps are displayed in the
Mollweide projection with a 130°E central meridian.
Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd
Dispersal barriers across the Indo-West Pacific
vs. annual), larval behaviour and competency characteristics,
reproductive output (fecundity and density of reproductive
adults), the pelagic larval duration (PLD) and larval mortal-
ity rate. Parameter values were informed by empirical data
from the literature (Table 1). Thus, our 99 hypothetical taxa
represent a broad range of biological traits, but these should
not be interpreted as being proportional to the relative
frequency of traits observed in nature.
Four generic larval behaviour strategies were modelled:
(1) Active strategy – actively swimming larvae within a
10 km ‘sensory halo’ of habitat and a 5-day pre-competency
period, (2) Passive strategy – passive larvae and a 5-day
pre-competency period, (3) Pelagic spawning – active swim-
ming larvae within the sensory zone and a 9-day pre-
competency period and (4) Benthic spawning – actively
swimming larvae within a sensory zone and a 1-day pre-
competency period. Reproductive output is likely to span
several orders of magnitude across species (Hughes et al.,
2000) and was modelled using an appropriate range
(103–107 larvae). Although the importance of PLD to
genetic connectivity is debatable (Weersing & Toonen, 2009;
Selkoe & Toonen, 2011), it may be important for long-dis-
tant dispersal (Bradbury et al., 2008; Faurby & Barber,
2012; Treml et al., 2012). Therefore, our simulations cov-
ered a range of maximum PLDs, from 10 to 60 days, repre-
senting the majority of marine taxa (Shanks, 2009; Mora
et al., 2012). Larval mortality has profound implications for
connectivity (Cowen et al., 2000), yet relatively little is
known regarding species-specific larval mortality in the
ocean. To accommodate this uncertainty, our study
included three levels of mortality: 10, 20 and 30% day 1,
broadly encompassing the known ranges for a variety of lar-
vae (Houde, 2002). Although these parameters represent a
diversity of marine taxa, several life histories are omitted,
such as those lacking a pelagic larval stage and those with
exceptionally long larval durations.
A single dispersal simulation consisted of modelling the
movement of a cloud of larvae from an individual reef patch
as it moves downstream, controlled by the biological param-
(J)
eters and ocean currents. When competent larvae encoun-
tered suitable habitat, these larvae settled and the density of
larvae was recorded at each time step. The model output was
Islands
(K) saved as a 3-dimensional matrix consisting of the discrete
dispersal kernels from every source population to every other
population, through time. These data were used to calculate
Vanuatu the population connectivity matrix representing the probabil-
ity that larvae released from each source (row) survive and
settle in each destination patch (column). See Treml et al.
1000 km (2012) for detailed numerical methods and sensitivity testing.
This connectivity matrix and the source population locations
(see Appendix S2) were used to construct georeferenced dis-
persal networks showing the geographic patterns of popula-
tion connectivity (Treml et al., 2008). The dispersal networks
for the 99 trait combinations were mapped to predict the
location and strength of ‘multitaxon’ dispersal barriers across
the IWP.
467
E. A. Treml et al.

Table 1 Ranges in parameter values for taxon-specific dispersal simulations. We varied spawning periodicity (seasonal vs. annual),
larval behaviour (active, passive, pelagic and benthic), pelagic larval duration (10–60 days as larvae), larval mortality (10, 20 and 30%
day 1) and relative reproductive output (103–107 larvae per population) to characterize 99 hypothetical model taxa. See Appendix S1 for
parameter sets for the 99 simulated taxa

Spawning Larval behaviour Relative reproductive Pelagic larval duration Larval mortality (% Unique model species (99
periodicity strategy output (days) day 1) total)

Annual Passive [103 105 107] [20 40 60] [10 20 30] 3 9 3 9 3 = 27

Annual Active [103 105 107] [20 40 60] [10 20 30] 3 9 3 9 3 = 27
Seasonal Pelagic spawner [103 105 107] [10 20 30] [10 20 30] 3 9 3 9 3 = 27
Seasonal Benthic spawner [103 105 107] [10 20] [10 20 30] 2 9 3 9 3 = 18

Table 2 Description of seascape barriers previously described for the region. The ID references the location of the barrier in Fig. 1. The
proportion of the 99 modelled taxa that cannot pass through the barrier was calculated. The percent of species filtered is quantified in
the direction implied in the description of the barrier, as well as in the reverse direction

Multispecies biophysical barriers Percent filtered

ID Barrier description Forward, % Reverse, % Barrier references

A Philippines to Taiwan 15.1 60.6 [1]

B North Equatorial Current, Philippines (S to N) 0.0 3.0 [2], [3]
C Micronesia to Guam 70.1 91.9 [4]
D Indo-Pacific to Palau 11.1 81.8 [1], [4], [5]
E Halmahera & Mindanao Eddies (E to W) 8.3 10.6 [6]
F Sunda Shelf/Java Sea (W to E) 17.0 22.6 [4], [7], [8]
G ‘Southern Barrier’ Java Sea (N to S) 4.0 9.4 [2], [6], [7], [8]
H Cenderawasih Bay to West 6.1 10.6 [6], [8], [9], [10]
I Cenderawasih Bay to East 17.5 19.5 [6], [11]
J Melanesia to Micronesia 24.8 83.7 [1]
K Solomons to Vanuatu 43.9 40.9 [4]
L Indonesia to Ningaloo, WA 71.7 92.8 [12], [13]

[1] Blum (1989); [2] Ravago-Gotanco et al. (2007); [3] Carpenter et al. (2011); [4] Veron et al. (2009); [5] Benzie & Williams (1995); [6] Barber
et al. (2006); [7] Barber et al. (2000); [8] Nuryanto & Kochzius (2009); [9] Crandall et al. (2008); [10] DeBoer et al. (2008); [11] Barber et al.
(2002); [12] Dethmers et al. (2006); [13] Imron et al. (2007).

To explore the strength of dispersal barriers, we calculated
the over-water distance between all 1002 reef habitat patches
and identified all potential connections falling within five
distance classes: 0–25 km, 25–50 km, 50–100 km, 100–
150 km and 150–200 km. These distance classes approximate
published thresholds for demographically significant popula-
tion connectivity from a variety of taxa and data sources
(Shanks et al., 2003; Siegel et al., 2003; Palumbi, 2004; Cowen
et al., 2006; Halpern et al., 2006; Bradbury et al., 2008;
Gaines et al., 2010). For all trait combinations, we calcu-
lated the proportion of these potential pathways that were
impermeable to the modelled taxa. Plotting the proportion
of potential connections that act as barriers in each distance
class and across all taxa illustrates the relationship between
over-water distance and seascape permeability.
To visualise and analyse the geographic configuration of
multitaxon dispersal barriers, we extracted a small subset of
the >1 million potential connections among the 1002 reef
patches. We chose 61 locations matching existing sampling
sites from published studies (e.g. Keyse et al., 2014) and
unpublished collections relating to marine population con-
nectivity (points in Fig. 1, labelled in Appendix S3). These
sites were then used to tessellate the seascape by drawing
Thiessen polygons around each sample site where boundaries
are equidistant from all nearest sites (Manni et al., 2004).
Each of the resultant 155 boundaries in this tessellation then
represents the approximate location of a potential biophysi-
cal barrier to be evaluated (in two directions) and formed
the basis for all subsequent analyses.
To quantify the relationship between the biological attri-
butes (Table 1) and the permeability of each of the 155
potential barriers, we used logistic regression and recursive
partitioning regression approaches. The main effects of all
biological and temporal parameters were quantified using the
logistic regression (binomial family, logit link function) for
two data matrices. The first assessed patterns across the 99
model taxa with the form: barrier permeability ~ PLD + mor-
tality + reproductive output + factor(Annual Active) + fac-
tor(Seasonal Benthic) + factor(Seasonal Pelagic). In this
regression, the reference level for the spawning strategy factor

468 Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd

was the Annual-spawning passive behaviour. The second
regression was used to quantify the effect of inter- and intra-
annual variability, in addition to the biological parameters,
within one behaviour class (seasonal spawning benthic behav-
iour) using the form: barrier permeability ~ PLD + mortal-
ity + reproductive output + factor(El Ni~ no) + factor(La Ni~na)
+ factor(Jan) + factor(Apr) + factor(Jul). In this regression,
the ‘neutral’ year and October month was used as the refer-
ence level for the factors. In both analyses, barrier permeabil-
ity was binary representing whether the potential barrier was
crossed (1) or not (0). Significant coefficients were then inter-
preted as their direct effect (strength and direction) on barrier
permeability across the seascape. To capture all significant
interactions among parameters, we used a variance-based
sensitivity analysis (Sobol, 2001) with a recursive partitioning
regression, implemented in R with the CompModSA package
(Storlie et al., 2009). The relative importance was quantified
using the total sensitivity index, T^j (Homma & Saltelli, 1996;
Storlie et al., 2009), which provides a single number summary
of the overall importance of each parameter on the response
variable, interpreted as the total proportion of the variability
that is due to each parameter, including all interactions with
other parameters (Storlie et al., 2009).
Finally, to map the barriers, we used the dispersal net-
works described above and intersected these with the Thies-
sen boundaries. We then calculated the proportion of taxa
that were unable to disperse between nearest neighbours of
the 61 sample sites (over multiple generations via stepping-
stones), producing a map of barriers weighted by the percent
of simulated taxa completely filtered at each boundary. All
previously described marine boundaries (Fig. 1) were evalu-
ated by extracting the coinciding Thiessen boundary. The
directional strength, in terms of the percent of taxa filtered,
was then calculated for each of the twelve previously
described barriers. For geographic context with respect to
multispecies connectivity, we summarized all dispersal con-
nections across all simulated taxa using a moving window
analysis to calculate the relative weighted density of connec-
tions within a 150-km radius. This line-density map reveals
the strength and spatial structure of multitaxa connectivity
throughout the IWP (Treml & Halpin, 2012).
RESULTS
Over 830,000 dispersal kernels were simulated across the
Indo-West Pacific for the 99 model taxa (data available in
Appendix S4). The dispersal networks for all taxa were used
to map the dispersal barriers across the region. Seascape per-
meability, revealed in the plot of barriers per distance class
(Fig. 2), decreases with increased over-water distance. For
the majority of modelled taxa, 5% of the over-water gaps
<25 km wide were impermeable, whereas over 40% of 150–
200 km gaps were impassable by most simulated taxa. In
addition to this decreasing permeability with increased dis-
tance, there was great variability across taxa within each dis-
tance class highlighting the strong influence of biology and
Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd
Dispersal barriers across the Indo-West Pacific
100
Proportion barriers (%)
60
20
0
25 50 100 150 200
Over−water distance (km)
Figure 2 Proportional barriers vs. over-water distance. Pairwise
over-water distances between all 1002 coral reef habitat patches
were measured, and the proportion of these potential linkages
that cannot be crossed by an individual taxon was determined.
All 99 taxa are summarized in each distance class. The number
of over-water gaps and the median value per distance bin are as
follows: <25 km (n = 864, ~x = 4.6%), 25–50 km (n = 1138,
~x = 9.4%), 50–100 km (n = 2250, ~x = 20.0%), 100–150 km
(n = 2758, ~x = 32.3%), 150–200 km (n = 3230, ~x = 41.0%).
geographic context on seascape permeability. Conversely, it
should be noted that across all distance classes the majority
of over-water gaps within the seascape can be crossed by the
majority of taxa explored (i.e. median values in all distance
classes were <50%).
The regression analyses quantify the main effects (bi coeffi-
cients in logistic regression) and the relative importance (T ^j
from sensitivity analysis using recursive partitioning regres-
sion) of each of three biological parameters (reproductive
output, pelagic larval duration and larval mortality) and tim-
ing (across years and months) on the permeability of each of
the potential barriers. The main effects derived from the
logistic regression analyses, across all potential boundaries,
were summarized for the 99 modelled taxa (Fig. 3a), and
separately for the benthic spawning strategy to explore inter-
and intraannual variability (Fig. 3b). Our main-effects logis-
tic regression model had significant explanatory power for
22% of the boundaries in the taxa-level analysis (R2 = 0.73).
Although a limited number of boundaries revealed a signifi-
cant relationship between permeability and these parameters,
the positive influence of reproductive output and PLD, and
the negative effect of larval mortality is clear (Fig. 3a). The
three behaviour and spawning strategies all showed an
increase in barrier permeability compared to the passive
behaviour reference level, with a stronger effect for the sea-
sonal spawning strategies. Exploring the full ensemble across
time for a single behavioural strategy (Fig. 3b) resulted in
90% of the boundaries with significant explanatory power
(mean adjusted R2 value of 0.63, median = 0.64). Reproduc-
tive output had the largest consistent effect, with a median
bi coefficient of 3.3, representing a 179 increase in the likeli-
hood of crossing potential barriers with an increase of one
order of magnitude in reproductive output. PLD had a med-
ian coefficient of 1.22, or a 49 increase in the likelihood of
crossing barriers with each 10 day increase in the larval
469
E. A. Treml et al.

(a)

20
15
Beta values
10
5
0
−5

Reproductive PLD Mortality Annual Seasonal Seasonal

Output Active Benthic Pelagic
Parameter

(b)
5

Figure 3 The main effects of biological

parameters on the permeability of
Beta values
0

potential dispersal barriers across the

seascape. (a) The logistic regression
coefficients, bi, resulting from the logistic
regression with 99 modelled taxa across
all potential barriers. (b) Logistic
−5

regression coefficients boxplot with the

324 dispersal simulations for a seasonal
spawning strategy and benthic behaviour
(3 years 9 4 seasons 9 3 PLDs 9 3
levels of mortality 9 3 levels of
−10

reproductive output). Only significant

coefficients (P > 0.05) from those
regressions resulting in a fit greater than
R2 = 0.50 are included (mean R2 = 0.65).
Reproductive PLD Mortality El Niño La Niña January April July The complementary recursive
Output
partitioning regression analysis is
Parameter presented in Appendix S5.

duration. Increasing larval mortality had a negative impact
on barrier permeability; an increase of 10% daily mortality
caused a 259 increase in not traversing boundaries. Years
(El Ni~
no and La Ni~ na compared to the ‘neutral’ year refer-
ence level) and months (Jan, Apr and Jul, compared to the
Oct reference level) all had highly variable effects on barrier
permeability.
Although the recursive partitioning regression analysis
included all main and interaction effects (see Appendix S5),
the patterns in parameter importance were highly consistent

470 Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd

with the logistic regression results. Similarly, the level of fit
across all regression models was high, with 89% having an
adjusted R2 value >0.50 (mean adjusted R2 = 0.74,
median = 0.78). The biological parameters displayed the
same rank order of importance (Reproductive Output >
PLD > Mortality), and the timing of larval release had mixed
importance across boundaries. Two additional bars were
added in Appendix S5, representing the cumulative impor-
tance across years and across months (Years, Months). These
cumulative importance values for spawning time, although
highly variable, were consistently ranked ahead of PLD in
terms of overall influence on determining the permeability of
boundaries.
The Thiessen polygon boundaries, derived from the 61
sample locations, were mapped showing the location and
strength of potential biophysical barriers across the seascape
(Fig. 4). The weighted relative density of all dispersal connec-
tions was plotted illustrating the range from highly dense and
strong connectivity (red) to sparse and weak or no connectiv-
ity (dark blue). Together, this map shows the regions of high
connectivity (dark red) as well as the configuration of dis-
persal barriers. Throughout the central IWP, irregular pat-
terns of connectivity are present, contiguous with many weak
barriers capable of filtering up to 10% of the modelled taxa.
Potential Barriers
% Taxa filtered
< 5%
5 to 10%
10 to 30%
30 to 50%
> 50%
1000 km
Figure 4 Multitaxon dispersal barriers and flow corridors. A
Thiessen tessellation algorithm was used to define boundaries
between all 61 sample sites. These sites and their shared
boundaries were used to quantify the proportion of simulated
taxa that cannot disperse between adjacent neighbours. The
weight of the edges in the tessellation represents the barrier’s
strength (proportion of taxa filtered). The coloured surface
shows the relative density of all potential dispersal connections
across all model taxa depicted with a linear histogram stretch
between the upper and lower 4th standard deviation. Warmest
colours are in areas of strong and dense connectivity.
Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd
Dispersal barriers across the Indo-West Pacific
Moving out of the central IWP, biophysical dispersal barriers
become stronger showing areas of significant dispersal limita-
tion for many taxa, with 30% to more than 50% of taxa
unable to pass to/from the central IWP. The proportion of
variance explained between the strength of biophysical barri-
ers and overwater or Euclidean distance was, at most 49%
(overwater R2 = 0.34, Euclidean distance R2 = 0.49). In other
words, over 50% of the variation in the strength of biophysi-
cal barriers is due to other biological or physical dynamics.
Using the location of twelve previously described marine
boundaries in the region (A–L, Fig. 1), we calculated the
proportion of modelled taxa that were unable to pass though
each of these putative barriers. Although directionality was
rarely implied in these previously described barriers, we have
reported the proportion of taxa filtered in both directions
for each location (Table 2). The proportion of modelled taxa
filtered ranged from 0% (barrier B) to 93% (barrier L), often
with strong directionality (e.g. barriers A, D and J). More
than 80% of the boundaries surveyed in this analysis were
capable of filtering more than 10% of the modelled taxa. In
general, those marine barriers described with respect to sis-
ter-species boundaries (e.g. Blum, 1989) and edges of species
ranges (e.g. Veron et al., 2009) showed a greater taxon-filter-
ing capacity (44–93% filtered taxa, barriers A, C, D, J, K, L)
than those described from within-species genetic differentia-
tion (0–23% filtered taxa, Table 2).
DISCUSSION
Identifying the location and strength of dispersal barriers has
been a great challenge in the marine environment and is
important for understanding biogeographic patterns, popula-
tion genetic structure, and metapopulation dynamics, as well
as providing the framework for developing regional conser-
vation plans. We have shown how life history characteristics,
based on empirical data, together with a regional ocean
model, can be used to predict barriers to dispersal. The pat-
terns we have identified are the result of dispersal dynamics
(biological traits and physical oceanography), suitable habitat
area and the spatial configuration of this habitat with respect
to ocean currents. Our results complement existing regional
research on biophysical dispersal patterns (Kool et al., 2011;
Wood et al., 2014), comparative phylogeography synthesis
(Crandall et al., 2008; Carpenter et al., 2011) and species-
specific population genetic studies by providing a comprehen-
sive regional picture of contemporary seascape permeability.
In addition, we suggest that the location and strength of mar-
ine dispersal barriers is largely determined by the reproductive
output, reproductive strategy (spawning time/frequency), and
the maximum length of the pelagic larval stage. These results
provide quantitative and spatially explicit hypotheses regard-
ing the location and strength of dispersal barriers, as well as a
regionally consistent framework for evaluating empirical esti-
mates of connectivity.
Although there are no empirical data covering the geo-
graphic and taxonomic scope of this study, we have shown
471
E. A. Treml et al.
broad support for many previously described marine bound-
aries (Table 2). For example, the connectivity hotspot and
surrounding barriers of the Halmahera region in western
PNG (E and H, Fig. 1 & Table 2) described in the genetic
analysis of several seastars (Crandall et al., 2008) and sto-
matopods (Barber et al., 2006), have the capacity to filter
10% of the modelled taxa and may represent formidable, yet
semi-permeable, dispersal barriers for a variety of species.
The Sunda Shelf (barrier F) has been recognized as a strong
barrier for several seahorses (Lourie & Vincent, 2004) and
giant clams (Nuryanto & Kochzius, 2009) and is shown here
to potentially filter up to 23% of the simulated taxa. The
location and strength of known species-level breaks are also
largely consistent with our results, especially between Palau
and the Indo-Pacific (barrier D), between the Solomon
Islands and Vanuatu (barrier K) and between the Philippines
and Taiwan (barrier A), as described for corals (Veron et al.,
2009) and butterflyfishes (Blum, 1989). The previously
described North Equatorial Current barrier in the Philippines
(Carpenter et al., 2011) was not strongly supported by this
analysis as it was predicted to filter up to at most 3% of the
simulated taxa (barrier B). Many other potential dispersal
barriers of varying strength have been identified where
empirical evidence is lacking (Fig. 4). In addition, a large
number of marine species are regarded as widespread
through the IWP, indicating the entire region is somewhat
permeable for a variety of taxa (Kulbicki et al., 2013). Our
results also support this observation in that many barriers
identified within the central IWP are quite permeable and fil-
ter, at most, 30% of the simulated taxa (Fig. 4). This strong
apparent duality in how the connectivity across the IWP is
perceived is evidence of the great biological–physical com-
plexity and is expected to remain the focus of much research
in the future.
Although the biophysical complexity in the IWP is a sig-
nificant predictor of species richness (Sanciangco et al.,
2013), the role of specific life histories in the development of
broad-scale biogeographic patterns has received relatively lit-
tle attention. Here, we have provided additional evidence
that three biological traits may significantly influence the
strength of barriers. Across all simulated taxa and all distance
classes, reproductive output consistently determined the per-
meability of seascape barriers (Fig. 3). This strong relation-
ship is consistent with the empirical work relating the levels
of recruitment with the production of larvae in corals at
large scales (Hughes et al., 2000). The large effect of spawn-
ing phenology, larval behaviour strategy, along with the max-
imum PLD, on seascape barrier permeability (Fig. 3) is also
supported by empirical and modelling studies (Huggett
et al., 2003; Shanks, 2009; Carson et al., 2010). The strong
yet variable effects of biological traits imply that a barrier, in
the strict sense of the word, is not required to disrupt move-
ment; habitat patches of low suitability or sink populations
may be sufficient.
In addition to mapping potential biogeographic breaks
throughout the Indo-West Pacific, these spatially explicit
472
dispersal predictions serve as testable hypotheses regarding
the role of dispersal in the development or maintenance of
these patterns. The dispersal networks (data in Appendices
S2 and S4) may be used as a proxy for ‘dispersal distance’
between sample sites, explicitly quantifying the potential
routes of connectivity. Of course potential dispersal corri-
dors and barriers are only one component in the complex
processes ultimately leading to the observed patterns in
genetic and community structure. Other important factors
include environmental variability and tolerances, historical
vicariance events, species interactions, density dependence,
survival following dispersal and other biophysical interac-
tions. Fortunately, several multispecies genetic databases are
currently being assembled through collaborative projects
and data sharing initiatives. Soon, we should be able to
test whether it is justifiable or appropriate to generalize
across taxa and identify community-level impacts resulting
from multispecies dispersal barriers such as the filtering of
species with increased isolation (e.g. Mora et al., 2003).
These comparative and cross-taxon studies could assist in
identifying the key biological traits and biophysical interac-
tions leading to these emergent patterns (Ayre et al.,
2009).
Several important caveats are required when interpreting
results presented here. (1) Although larval retention often
dominates local-scale connectivity patterns, these values were
removed from the analysis as they do not contribute to
broad-scale connectivity, other than reducing the relative
strength of downstream connections. (2) The spatio-tempo-
ral resolution and extent of the modelled hydrodynamic data
are limiting. Although in using three specific years with
respect to the El Ni~ no Southern Oscillation cycle (1997,
1999, 2001), we captured a range in ocean dynamics includ-
ing seasonal, decadal, and monsoonal patterns, modelling
across many more years would be advantageous. Similarly,
fine-scale hydrodynamics such as tides are rarely included in
regional ocean models, and as a result, local-scale connectiv-
ity patterns may be underestimated. Nonetheless, we believe
the impact on our seascape-level predictions would be
minor. Underestimating local retention would lead to an
overestimation of downstream connectivity and therefore
would result in a conservative estimate of barrier strength –
an accepted compromise in our analysis. (3) There may be
significant discrepancies between the time-scale of the dis-
persal simulations and the time-scales inherent in empirical
data (e.g. genetic data summarizing ecological processes over
many thousands of generations). Nonetheless, our results
provide spatially explicit and quantitative predictions that
can then be used to better investigate biogeographic and
genetic patterns. (4) Other than habitat area, all other habitat
patch characteristics were assumed to be identical, which
may impact the predictions of realized dispersal patterns. In
the future, including environmental differences such as
temperature, salinity and quality, and species’ tolerances
would no doubt improve the predictions of realized dispersal
barriers.
Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd

CONCLUSION
Combining biophysical models of larval dispersal within a
spatially explicit framework allows the geographic patterns of
dispersal barriers to be mapped. Exploring the relationship
between barrier formation and biological characteristics
revealed the importance of reproductive output, reproductive
phenology and pelagic larval duration – each worthy of
future empirical investigation. We found a 109 increase in
reproductive output increased the chance of crossing dis-
persal barriers by 179, a 10 day increase in PLD increased
the odds of crossing by four times and a 10% increase in lar-
val mortality resulted in a 259 decrease in the odds of cross-
ing marine barriers. This biological and physical complexity
resulted in largely intuitive geographic pattern in barriers,
with weak barriers common within the central Indo-West
Pacific and strong multispecies barriers at the periphery. We
posit that this approach can contribute to the development
of an accurate bioregionalization and robust conservation
prioritization across the Indo-West Pacific, and fill gaps in
our understanding of the biogeography. We hope the
approach presented here provides a useful framework for
identifying and testing the effects of dispersal barriers in
marine ecosystems.
ACKNOWLEDGEMENTS
We thank Steve Swearer and Sean Connolly and three anon-
ymous referees for their constructive comments that greatly
improved the manuscript. Funding for this work was pro-
vided by the Australian Research Council (DP0878306 to CR
and HP), the World Wildlife Fund (Kathryn Fuller Post-doc-
toral Research Fellowship to EAT) and the National Science
Foundation (NSF OISE–0730256 to PNH).
REFERENCES
Ayre, D.J., Minchinton, T.E. & Perrin, C. (2009) Does life
history predict past and current connectivity for rocky
intertidal invertebrates across a marine biogeographic bar-
rier? Molecular Ecology, 18, 1887–1903.
Barber, P.H., Palumbi, S.R., Erdmann, M.V. & Moosa, M.K.
(2000) Biogeography – a marine Wallace’s line? Nature,
406, 692–693.
Barber, P.H., Palumbi, S.R., Erdmann, M.V. & Moosa, M.K.
(2002) Sharp genetic breaks among populations of Hap-
tosquilla pulchella (Stomatopoda) indicate limits to larval
transport: patterns, causes, and consequences. Molecular
Ecology, 11, 659–674.
Barber, P.H., Erdmann, M.V. & Palumbi, S.R. (2006) Com-
parative phylogeography of three codistributed stomato-
pods: origins and timing of regional lineage diversification
in the coral triangle. Evolution, 60, 1825–1839.
Baums, I.B., Miller, M.W. & Hellberg, M.E. (2005)
Regionally isolated populations of an imperiled Caribbean
coral, Acropora palmata. Molecular Ecology, 14, 1377–1390.
Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd
Dispersal barriers across the Indo-West Pacific
Benzie, J.A.H. & Williams, S.T. (1995) Gene flow among
giant clam (Tridacna gigas) populations in Pacific does not
parallel ocean circulation. Marine Biology, 123, 781–787.
Blum, S. (1989) Biogeography of the chaetodontidae: an
analysis of allopatry among closely related species. Environ-
mental Biology of Fishes, 25, 9–31.
Bowen, B.W., Rocha, L.A., Toonen, R.J. & Karl, S.A. (2013)
The origins of tropical marine biodiversity. Trends in Ecol-
ogy & Evolution, 28, 359–366.
Bradbury, I.R., Laurel, B., Snelgrove, P.V.R., Bentzen, P. &
Campana, S.E. (2008) Global patterns in marine dispersal
estimates: the influence of geography, taxonomic category
and life history. Proceedings of the Royal Society of London
Series B, Biological Sciences, 275, 1803–1809.
Burke, L., Reytar, K., Spalding, M.D. & Perry, A. (2012) Reefs
At Risk Revisited in the Coral Triangle, pp. 130. World
Resources Institute, Washington, DC.
Carpenter, K.E., Barber, P.H., Crandall, E.D., Ablan-Lagman,
M.C.A., Ambariyanto, Mahardika, G.N., Manjaji-Matsum-
oto, B.B., Juinio-Menez, M.A., Santos, M.D., Starger, C.J.
& Toha, A.H.A. (2011) Comparative phylogeography of
the coral triangle and implications for marine manage-
ment. Journal of Marine Biology, 2011, 1–14.
Carson, H.S., L opez-Duarte, P.C., Rasmussen, L., Wang, D.
& Levin, L.A. (2010) Reproductive timing alters population
connectivity in marine metapopulations. Current Biology,
20, 1926–1931.
Cowen, R.K. & Sponaugle, S. (2009) Larval dispersal and
marine population connectivity. Annual Review of Marine
Science, 1, 443–466.
Cowen, R.K., Lwiza, K.M.M., Sponaugle, S., Paris, C.B. & Ol-
son, D.B. (2000) Connectivity of marine populations: open
or closed? Science, 287, 857–859.
Cowen, R.K., Paris, C.B. & Srinivasan, A. (2006) Scaling
of connectivity in marine populations. Science, 311, 522–
527.
Crandall, E.D., Frey, M.A., Grosberg, R.K. & Barber, P.H.
(2008) Contrasting demographic history and phylogeo-
graphical patterns in two Indo-Pacific gastropods. Molecu-
lar Ecology, 17, 611–626.
Crandall, E.D., Treml, E.A., Liggins, L., Gleeson, L., Yasuda,
N., Barber, P.H., Worheide, G. & Riginos, C. (2014)
Return of the ghosts of dispersal past: historical spread and
contemporary gene flow in the blue sea star Linckia laevig-
ata. Bulletin of Marine Science, 90, 399–425.
Davies, S.W., Treml, E.A., Kenkel, C.D. & Matz, M.V. (2015)
Exploring the role of Micronesian islands in the mainte-
nance of coral genetic diversity in the Pacific Ocean.
Molecular Ecology, 24, 70–82.
DeBoer, T., Subia, M., Erdmann, M., Kovitvongsa, K. & Bar-
ber, P.H. (2008) Phylogeography and limited genetic con-
nectivity in the endangered boring giant clam across the
Coral Triangle. Conservation Biology, 22, 1255–1266.
Dethmers, K.E.M., Broderick, D., Moritz, C., Fitzsimmons,
N.N., Limpus, C.J., Lavery, S., Whiting, S., Guinea, M.,
Prince, R.I.T. & Kennett, R. (2006) The genetic structure
473
E. A. Treml et al.
of Australasian green turtles (Chelonia mydas): exploring
the geographical scale of genetic exchange. Molecular Ecol-
ogy, 15, 3931–3946.
Faurby, S. & Barber, P.H. (2012) Theoretical limits to the
correlation between pelagic larval duration and population
genetic structure. Molecular Ecology, 21, 3419–3432.
Gaines, S.D., Lester, S.E., Grorud-Colvert, K., Costello, C. &
Pollnac, R. (2010) Evolving science of marine reserves: new
developments and emerging research frontiers. Proceedings
of the National Academy of Sciences USA, 107, 18251–
18255.
Gaither, M.R. & Rocha, L.A. (2013) Origins of species rich-
ness in the Indo-Malay-Philippine biodiversity hotspot:
evidence for the centre of overlap hypothesis. Journal of
Biogeography, 40, 1638–1648.
Galindo, H.M., Olson, D.B. & Palumbi, S.R. (2006) Seascape
Genetics: a coupled oceanographic-genetic model predicts
population structure of Caribbean corals. Current Biology,
16, 1622–1626.
Halpern, B.S., Regan, H.M., Possingham, H.P. & McCarthy,
M.A. (2006) Accounting for uncertainty in marine reserve
design. Ecology Letters, 9, 2–11.
Hastings, A. & Botsford, L.W. (2006) A simple persistence
condition for structured populations. Ecology Letters, 9,
846–852.
Hickerson, M.J. & Meyer, C.P. (2008) Testing comparative
phylogeographic models of marine vicariance and dispersal
using a hierarchical Bayesian approach. BMC Evolutionary
Biology, 8, 18.
Hobbs, J.-P., Jones, G. & Munday, P. (2010) Rarity and
extinction risk in coral reef angelfishes on isolated islands:
interrelationships among abundance, geographic range size
and specialisation. Coral Reefs, 29, 1–11.
Hoeksema, B.W. (2007) Delineation of the Indo-Malayan
centre of maximum marine biodiversity: The coral triangle.
Topics in Geobiology (ed. by W. Renema), pp. 117–178.
Springer, Dordrecht, Netherlands.
Homma, T. & Saltelli, A. (1996) Importance measures in
global sensitivity analysis of nonlinear models. Reliability
Engineering & System Safety, 52, 1–17.
Houde, E.D. (2002) Mortality. Fishery Science: The Unique
Contributions of Early Life Stages (ed. by L.A. Fuiman and
R.G. Werner), pp. 64–87. Blackwell Publishing, Oxford.
Huggett, J., Freon, P., Mullon, C. & Penven, P. (2003) Mod-
elling the transport success of anchovy Engraulis encrasico-
lus eggs and larvae in the southern Benguela: the effect of
spatio-temporal spawning patterns. Marine Ecology Progress
Series, 250, 247–262.
Hughes, T.P., Baird, A.H., Dinsdale, E.A., Moltschaniwskyj,
N.A., Pratchett, M.S., Tanner, J.E. & Willis, B.L. (2000)
Supply-side ecology works both ways: the link between
benthic adults, fecundity, and larval recruits. Ecology, 81,
2241–2249.
Imron, Jeffrey, B., Hale, P., Degnan, B.M. & Degnan, S.M.
(2007) Pleistocene isolation and recent gene flow in Halio-
474
tis asinina, an Indo-Pacific vetigastropod with limited dis-
persal capacity. Molecular Ecology, 16, 289–304.
Jones, G.P., Srinivasan, A. & Almany, G.R. (2007) Population
connectivity and conservation of marine biodiversity.
Oceanography, Special Issue on Marine Population Connec-
tivity, 20, 100–111.
Keyse, J., Crandall, E.D., Toonen, R.J., Meyer, C.P., Treml,
E.A. & Riginos, C. (2014) The scope of published popula-
tion genetic data for Indo-Pacific marine fauna and future
research opportunities in the region. Bulletin of Marine Sci-
ence, 90, 47–78.
Kinlan, B.P. & Gaines, S.D. (2003) Propagule dispersal in
marine and terrestrial environments: a community perspec-
tive. Ecology, 84, 2007–2020.
Knowlton, N. & Weigt, L.A. (1998) New dates and new rates
for divergence across the Isthmus of Panama. Proceedings
of the Royal Society of London Series B, Biological Sciences,
265, 2257–2263.
Kool, J.T., Paris, C.B., Barber, P.H. & Cowen, R.K. (2011)
Connectivity and the development of population genetic
structure in Indo-West Pacific coral reef communities. Glo-
bal Ecology and Biogeography, 20, 695–706.
Kulbicki, M., Parravicini, V., Bellwood, D.R., Arias-Gonzalez,
E., Chabanet, P., Floeter, S.R., Friedlander, A., McPherson,
J., Myers, R.E., Vigliola, L. & Mouillot, D. (2013) Global
biogeography of reef fishes: a hierarchical quantitative
delineation of regions. PLoS ONE, 8, e81847.
Largier, J.L. (2003) Considerations in estimating larval dis-
persal distances from oceanographic data. Ecological Appli-
cations, 13, S71–S89.
Lessios, H.A. & Robertson, D.R. (2006) Crossing the impass-
able: genetic connections in 20 reef fishes across the eastern
Pacific barrier. Proceedings of the Royal Society of London
Series B, Biological Sciences, 273, 2201–2208.
Lourie, S.A. & Vincent, A.C.J. (2004) Using biogeography to
help set priorities in marine conservation. Conservation
Biology, 18, 1004–1020.
MacArthur, R. & Wilson, E. (1967) The Theory of Island Bio-
geography. Princeton University Press, Princeton, NJ.
Manni, F., Guerard, E. & Heyer, E. (2004) Geographic pat-
terns of (genetic, morphologic, linguistic) variation: how
barriers can be detected by using Monmonier’s algorithm.
Human Biology, 76, 173–190.
Marshall, D.J., Monro, K., Bode, M., Keough, M.J. &
Swearer, S. (2010) Phenotype-environment mismatches
reduce connectivity in the sea. Ecology Letters, 13, 128–
140.
Mayr, E. (1963) Animal Species and Evolution. Harvard Uni-
versity Press, Cambridge, MA.
Meyer, C.P., Geller, J.B. & Paulay, G. (2005) Fine scale
endemism on coral reefs: archipelagic differentiation in
turbinid gastropods. Evolution, 59, 113–125.
Mirams, A.G.K., Treml, E.A., Shields, J.L., Liggins, L. & Rigi-
nos, C. (2011) Vicariance and dispersal across an intermit-
tent barrier: population genetic structure of marine
Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd

animals across the Torres Strait land bridge. Coral Reefs,
30, 937–949.
Mora, C., Chittaro, P.M., Sale, P.F., Kritzer, J.P. & Ludsin,
S.A. (2003) Patterns and processes in reef fish diversity.
Nature, 421, 933–936.
Mora, C., Treml, E.A., Roberts, J., Crosby, K., Roy, D. & Tit-
tensor, D.P. (2012) High connectivity among habitats pre-
cludes the relationship between dispersal and range size in
tropical reef fishes. Ecography, 35, 89–96.
North, E.W., Gallego, A. & Petitgas, P. (2009) Manual of rec-
ommended practices for modelling physical - biological
interactions during fish early life. ICES Cooperative
Research Report, 295, i–iii, 1-112.
Nuryanto, A. & Kochzius, M. (2009) Highly restricted gene
flow and deep evolutionary lineages in the giant clam Trid-
acna maxima. Coral Reefs, 28, 607–619.
Palumbi, S.R. (1994) Genetic-divergence, reproductive isola-
tion, and marine speciation. Annual Review of Ecology and
Systematics, 25, 547–572.
Palumbi, S.R. (2004) Marine reserves and ocean neighbor-
hoods: the spatial scale of marine populations and their
management. Annual Review of Environment and Resources,
29, 31–68.
Quinn, T.M., Taylor, F.W. & Crowley, T.J. (1993) A 173 year
stable-isotope record from a tropical south-pacific coral.
Quaternary Science Reviews, 12, 407–418.
Randall, J.E. (1998) Zoogeography of shore fishes of the
Indo-Pacific region. Zoological Studies, 37, 227–268.
Ravago-Gotanco, R.G., Magsino, R.M. & Juinio-Menez, M.A.
(2007) Influence of the North Equatorial Current on the
population genetic structure of Tridacna crocea (Mollusca:
Tridacnidae) along the eastern Philippine seaboard. Marine
Ecology Progress Series, 336, 161–168.
Riginos, C. & Liggins, L. (2013) Seascape Genetics: popula-
tions, individuals, and genes marooned and adrift. Geogra-
phy Compass, 7, 197–216.
Riginos, C., Douglas, K.E., Jin, Y., Shanahan, D.F. & Treml,
E.A. (2011) Effects of geography and life history traits on
genetic differentiation in benthic marine fishes. Ecography,
34, 566–575.
Roberts, C.M., McClean, C.J., Veron, J.E.N., Hawkins,
J.P., Allen, G.R., McAllister, D.E., Mittermeier, C.G.,
Schueler, F.W., Spalding, M., Wells, F., Vynne, C. &
Werner, T.B. (2002) Marine biodiversity hotspots and
conservation priorities for tropical reefs. Science, 295,
1280–1284.
Sanciangco, J.C., Carpenter, K.E., Etnoyer, P.J. & Moretz-
sohn, F. (2013) Habitat availability and heterogeneity and
the indo-pacific warm pool as predictors of marine spe-
cies richness in the tropical indo-pacific. PLoS ONE, 8,
e56245.
Sandin, S.A., Vermeij, M.J.A. & Hurlbert, A.H. (2008) Island
biogeography of Caribbean coral reef fish. Global Ecology
and Biogeography, 17, 770–777.
Selkoe, K.A. & Toonen, R.J. (2011) Marine connectivity: a
new look at pelagic larval duration and genetic metrics
Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd
Dispersal barriers across the Indo-West Pacific
of dispersal. Marine Ecology Progress Series, 436, 291–
305.
Shanks, A.L. (2009) Pelagic larval duration and dispersal dis-
tance revisited. Biological Bulletin, 216, 373–385.
Shanks, A.L., Grantham, B.A. & Carr, M.H. (2003) Propagule
dispersal distance and the size and spacing of marine
reserves. Ecological Applications, 13, S159–S169.
Siegel, D.A., Kinlan, B.P., Gaylord, B. & Gaines, S.D. (2003)
Lagrangian descriptions of marine larval dispersion. Marine
Ecology-Progress Series, 260, 83–96.
Slatkin, M. (1987) Gene flow and the geographic structure of
natural-populations. Science, 236, 787–792.
Sobol, I.M. (2001) Global sensitivity indices for nonlinear
mathematical models and their Monte Carlo estimates.
Mathematics and Computers in Simulation, 55, 271–280.
Spalding, M., Ravilious, C. & Green, E.P. (2001) World Atlas
of Coral Reefs. University of California Press, Berkely.
Steneck, R., Paris, C., Arnold, S., Ablan-Lagman, M., Alcala,
A., Butler, M., McCook, L., Russ, G. & Sale, P. (2009)
Thinking and managing outside the box: coalescing con-
nectivity networks to build region-wide resilience in coral
reef ecosystems. Coral Reefs, 28, 367–378.
Storlie, C.B., Swiler, L.P., Helton, J.C. & Sallaberry, C.J.
(2009) Implementation and evaluation of nonparametric
regression procedures for sensitivity analysis of computa-
tionally demanding models. Reliability Engineering & Sys-
tem Safety, 94, 1735–1763.
Timm, J. & Kochzius, M. (2008) Geological history and
oceanography of the Indo-Malay Archipelago shape the
genetic population structure in the false clown anemone-
fish (Amphiprion ocellaris). Molecular Ecology, 17, 3999–
4014.
Treml, E.A. & Halpin, P.N. (2012) Marine population con-
nectivity identifies ecological neighbors for conservation
planning in the Coral Triangle. Conservation Letters, 5,
441–449.
Treml, E., Halpin, P., Urban, D. & Pratson, L. (2008) Model-
ing population connectivity by ocean currents, a graph-the-
oretic approach for marine conservation. Landscape
Ecology, 23, 19–36.
Treml, E.A., Roberts, J.J., Chao, Y., Halpin, P.N., Possing-
ham, H.P. & Riginos, C. (2012) Reproductive output and
duration of the pelagic larval stage determine seascape-wide
connectivity of marine populations. Integrative and Com-
parative Biology, 52, 525–537.
Veron, J.E.N., Devantier, L.M., Turak, E., Green, A.L., Kinin-
month, S., Stafford-Smith, M. & Peterson, N. (2009)
Delineating the Coral Triangle. Galaxea Journal of Coral
Reef Studies, 11, 91–100.
Wang, X. & Chao, Y. (2004) Simulated Sea Surface Salinity
variability in the tropical Pacific. Geophysical Research Let-
ters, 31, L02302.
Wang, P., Song, Y.T., Chao, Y. & Zhang, H.C. (2005) Parallel
computation of the regional ocean modeling system. Inter-
national Journal of High Performance Computing Applica-
tions, 19, 375–385.
475
E. A. Treml et al.
Weersing, K. & Toonen, R.J. (2009) Population genetics, lar-
val dispersal, and connectivity in marine systems. Marine
Ecology Progress Series, 393, 1–12.
Werner, F.E., Cowen, R.K. & Paris, C.B. (2007) Coupled bio-
logical and physical models: present capabilities and neces-
sary developments for future studies of population
connectivity. Oceanography, Special Issue on Marine Popula-
tion Connectivity, 20, 54–69.
White, C., Selkoe, K.A., Watson, J., Siegel, D.A., Zacherl,
D.C. & Toonen, R.J. (2010) Ocean currents help explain
population genetic structure. Proceedings of the Royal
Society of London Series B, Biological Sciences, 277, 1685–
1694.
Wood, S., Paris, C.B., Ridgwell, A. & Hendy, E.J. (2014)
Modelling dispersal and connectivity of broadcast spawning
corals at the global scale. Global Ecology and Biogeography,
23, 1–11.
SUPPORTING INFORMATION
Additional Supporting Information may be found in the
online version of this article:
Appendix S1 Parameter sets for the 99 model taxa, for all
combinations.
Appendix S2 List of reef IDs and centroid locations.
Appendix S3 Sample locations and labels.
476
Appendix S4 Row-normalized probability matrices for the
99 trait combinations as a three-dimensional array for read-
ing in the R software environment. Array indexing as [from
reef, to reef, model taxon].
Appendix S5 Recursive partitioning figures.
DATA ACCESSIBILITY
All reef data, connectivity matrices (Appendix S2 and S4)
and an example R script are publicly available from the
Dryad Digital Repository: http://dx.doi.org/10.5061/
dryad.24tt2
BIOSKETCH
Eric A. Treml is a lecturer in marine biology in the School
of BioSciences at the University of Melbourne. His research
is focused primarily on the spatial ecology of marine systems,
testing hypotheses regarding marine population connectivity
and exploring the implications for marine conservation and
ecosystem-based management. His research integrates
dynamic modelling, spatial analysis, graph theory and popu-
lation genetic techniques.
Editor: Robert Cowie
Diversity and Distributions, 21, 465–476, ª 2015 John Wiley & Sons Ltd