Tropical seamounts as stepping-stones for coral reef fishes: range extensions and new regional distributions from mesophotic ecosystems in the Coral Sea, Australia

Abstract

Seamounts and remote oceanic islands serve as valuable natural laboratories in which to study patterns and processes in marine biodiversity. A central hypothesis arising from studies of these systems is the ecological function of seamounts as stepping-stones for dispersal and population connectivity. Evidence of this mechanism exists for a range of taxa, including coral reef fishes, but is still lacking from many tropical seamounts in remote regions. In this study, we used remotely operated vehicles and baited remote underwater video systems to survey fish and benthic communities between 1 and 100 m on seamounts in the Coral Sea Marine Park (CSMP), Australia. We found evidence to support the stepping-stone model of ecological connectivity from new observations of 16 coral reef fishes which have previously not been recorded by quantitative surveys in the region. The widespread distribution of many of these species throughout the full latitudinal extent of the CSMP suggests that there is greater connectivity between mesophotic habitats in the Coral Sea and surrounding biogeographic regions than previously known. We also found a wide variety of mesophotic habitats and recorded significant depth range extensions for 78 fishes in these habitats. This further highlights the potential role of increased habitat area and heterogeneity in a stepping-stone effect throughout the region. Four of the fish occurrence records represent significant range extensions into the Coral Sea from adjacent biogeographic regions, and 13 fishes recorded by this study in the CSMP are not known from the neighbouring Great Barrier Reef, despite its close proximity. Although the Coral Sea remains relatively understudied, these findings suggest that larger-scale models of marine biogeography are relevant to communities in the region, particularly at mesophotic depths. Given the extent and the spatial arrangement of seamounts in the Coral Sea, our findings emphasise that the region is an important link between the centre of marine biodiversity in the Coral Triangle and the Southwest Pacific. Greater mesophotic sampling effort and genetic studies are necessary to understand the nature of connectivity and to establish the role of regional seamount chains, like the Coral Sea reefs, in broader marine biogeographic processes.

Full text

Vol.:(0123456789)
Marine Biodiversity (2024) 54:17
https://doi.org/10.1007/s12526-024-01404-0
ORIGINAL PAPER
Tropical seamounts asstepping‑stones forcoral reef fishes: range
extensions andnew regional distributions frommesophotic
ecosystems intheCoral Sea, Australia
G.F.Galbraith1,2 · B.J.Cresswell1,2 · E.C.McClure1,2 · A.S.Hoey1,2
Received: 23 April 2023 / Revised: 30 December 2023 / Accepted: 11 January 2024
© The Author(s) 2024
Abstract
Seamounts and remote oceanic islands serve as valuable natural laboratories in which to study patterns and processes in
marine biodiversity. A central hypothesis arising from studies of these systems is the ecological function of seamounts as
stepping-stones for dispersal and population connectivity. Evidence of this mechanism exists for a range of taxa, including
coral reef fishes, but is still lacking from many tropical seamounts in remote regions. In this study, we used remotely oper-
ated vehicles and baited remote underwater video systems to survey fish and benthic communities between 1 and 100 m
on seamounts in the Coral Sea Marine Park (CSMP), Australia. We found evidence to support the stepping-stone model of
ecological connectivity from new observations of 16 coral reef fishes which have previously not been recorded by quantitative
surveys in the region. The widespread distribution of many of these species throughout the full latitudinal extent of the CSMP
suggests that there is greater connectivity between mesophotic habitats in the Coral Sea and surrounding biogeographic
regions than previously known. We also found a wide variety of mesophotic habitats and recorded significant depth range
extensions for 78 fishes in these habitats. This further highlights the potential role of increased habitat area and heterogeneity
in a stepping-stone effect throughout the region. Four of the fish occurrence records represent significant range extensions into
the Coral Sea from adjacent biogeographic regions, and 13 fishes recorded by this study in the CSMP are not known from the
neighbouring Great Barrier Reef, despite its close proximity. Although the Coral Sea remains relatively understudied, these
findings suggest that larger-scale models of marine biogeography are relevant to communities in the region, particularly at
mesophotic depths. Given the extent and the spatial arrangement of seamounts in the Coral Sea, our findings emphasise that
the region is an important link between the centre of marine biodiversity in the Coral Triangle and the Southwest Pacific.
Greater mesophotic sampling effort and genetic studies are necessary to understand the nature of connectivity and to establish
the role of regional seamount chains, like the Coral Sea reefs, in broader marine biogeographic processes.
Keywords Marine biodiversity· ROVs· BRUVs· Biogeography· Patch habitats
Introduction
The Coral Sea, in the southwest Pacific Ocean, is the
second largest tropical marginal sea on earth and is
characterised by a complex bathymetry and diverse
seascape of distinct marine habitats (Ceccarelli etal.
2013; McKinnon etal. 2014). These habitats include
the deep sea, submerged banks, canyons, island chains
and large oceanic coral reef systems atop of seamounts
rising from deep waters (up to 3000 m) (Davies etal.
1989; Bridge etal. 2019). Ecologically, seamounts have
been variously considered as either stepping-stones for
marine dispersal that can promote regional connectivity
via chains of suitable habitat across deep open oceans
This article is a contribution to the Topical Collection Seamounts
and oceanic archipelagos and their role for the biodiversity,
biogeography, and dispersal of marine organisms.
Communicated by K. H. George
* G. F. Galbraith
gemma.galbraith@jcu.edu.au
1 Marine Biology andAquaculture, College ofScience
andEngineering, James Cook University, Townsville,
Queensland4811, Australia
2 ARC Centre ofExcellence forCoral Reef Studies, James
Cook University, Townsville, Queensland4811, Australia

Marine Biodiversity (2024) 54:17 17 Page 2 of 18
(Hubbs 1959; Rowden etal. 2010; Mazzei etal. 2021)
and conversely, as isolated islands that can give rise to
unique ecological communities (Richer de Forges etal.
2000; Hobbs etal. 2008, 2012; McClain etal. 2009).
The general application of these contrasting hypotheses
remains unclear and it is broadly agreed that the ecological
function of seamounts in marine connectivity depends on
multiple factors including geomorphology, depth, spatial
isolation, oceanographic processes and taxa-specific
dispersal capabilities (Mcclain 2007; Clark and Bowden
2015; Miller and Gunasekera 2017; Pinheiro etal. 2017).
The basis of the function of seamounts as stepping-stones
for the dispersal of marine organisms depends on the avail-
ability of suitable habitat at a given seamount, the distance
between seamounts and the nature and direction of ocean
currents and localised seamount-generated flows (Rowden
etal. 2010). The typical arrangement of seamounts in linear
chains is a particularly important feature of this hypothesis.
In combination with large-scale ocean currents, this spatial
arrangement can facilitate the successive transport of marine
larvae across oceanic basins or extending from mainland
coastlines (Mazzie etal. 2021; Simon etal. 2022). Moreover,
localised seamount-generated hydrodynamics are a unique
feature of these habitats and the stepping-stone model of
connectivity. These mechanisms may include closed recircu-
lating currents that can trap and retain larvae over seamount
summits, subsequently enhancing recruitment and settle-
ment along seamount chains (Mulineaux and Mills 1997;
Sponaugle etal. 2002). Despite wide recognition as highly
productive biodiversity hotspots and this important function
for connectivity in both tropical and temperate oceans, sea-
mounts remain one of the least explored and studied marine
biomes on earth (Clark etal. 2010b; Wagner etal. 2020;
Yesson etal. 2021). Consequently, many regional seamount
chains require considerably greater sampling effort to estab-
lish patterns of biodiversity and the mechanisms driving
ecological connectivity with wider biogeographic regions
(Rogers 2018).
Most seamount reef systems in the Coral Sea occur in
Australia’s Exclusive Economic Zone (EEZ) and are man-
aged as the Coral Sea Marine Park (CSMP). Together with
the French Natural Park of the Coral Sea (Le Parc Naturel de
la Mer de Corail), the Coral Sea possesses the largest com-
bined protected area in the world (Director of National Parks
2018). In Australia’s CSMP, over 30 individual reef systems
are spread across 22° of latitude on the Queensland and Mar-
ion Plateaus and constitute ~24,000 km2 of shallow-water (<
30 m) emergent coral reef habitat (Bridge etal. 2019). The
CSMP is bordered by major global marine biodiversity and
productivity hotspots; Australia’s Great Barrier Reef (GBR)
to the west, the Coral Triangle (specifically, Papua New
Guinea and the Solomon Islands) to the north, Vanuatu and
New Caledonia to the east and the Tasman Sea to the south
(Fig.1). It is therefore not surprising that the CSMP sup-
ports a relatively high diversity of reef fish (~1200 species)
and high abundance and biomass of sharks and other large
predatory fishes (Randall etal.1997; Ceccarelli etal. 2013;
Stuart-Smith etal. 2013; Hoey etal. 2022). Shallow-water
reef habitats in the CSMP have also been shown to sup-
port unique coral and reef fish communities that are distinct
from those of the adjacent GBR to the west and share more
similarities with those found in New Caledonia to the east
(Ceccarelli etal. 2013; Hoey etal. 2020). Oceanographic
processes and historic environmental conditions explain a
significant proportion of the evolutionary processes driving
genetic connectivity and biodiversity patterns within Coral
Sea populations and between surrounding regions (Cec-
carelli etal. 2013; Kessler and Cravatte 2013; Payet etal.
2022). However, the extent and spatial arrangement of coral
reef habitat on seamounts in the Coral Sea is also a signifi-
cant component of ecological connectivity in this region and
within the wider Central Pacific Ocean.
Although recent large-scale monitoring efforts (Hoey
etal. 2020, 2022) and some baseline surveys (Ayling
and Ayling 1984; Oxley etal. 2004) have established
quantitative ecological data for shallow-water coral reefs
in the CSMP, the remote nature of reefs in the Coral Sea
mean, they remain poorly documented compared to those
in the surrounding GBR or the Coral Triangle (Ceccarelli
2010). Additionally, there is a paucity of research
conducted on coral reefs below 30 m compared to shallow
reefs, particularly in Australia (Pyle and Copus 2019,
Eyal etal. 2021). Mesophotic coral ecosystems (MCEs)
are defined as light-dependent coral reef communities in
depths of 30–150 m (Loya etal. 2016). The exceptionally
clear oligotrophic waters of the Coral Sea allow light
penetration to considerable depths,and several exploratory
studies have confirmed the presence of MCEs in the
region to depths of up to 125m (Sarano and Pichon 1988;
Bongaerts etal. 2011; Muir etal. 2015; Englebert etal.
2015, 2017). At these depths, the complex bathymetries
of the Coral Sea seamounts are also highly variable, both
among and within individual reef systems (Harris etal.
2003; Beaman 2012). Some rise as vertical walls to the
surface (e.g. Bougainville and Osprey reefs), while others
have less abrupt slopes and many possess near-horizontal
areas along flanks and submerged shelves (e.g. Holmes
Reefs and East Diamond Islet). On more moderate slopes,
where hard substrate is present and light availability is
optimal, mesophotic depths can support high percentage
cover of photosynthetic habitat-forming taxa (Pérez-
Rosales etal. 2022). Hard and soft corals, macroalgae,
sponges and large benthic foraminifera are all important
constituents of MCEs and in turn provide habitats for other
marine organisms (Slattery and Lesser 2012; Lesser etal.
2018). This said, the extent of MCEs or other important

Marine Biodiversity (2024) 54:17 Page 3 of 18 17
benthic habitats between 0 and 150 m have not been
quantified or confirmed at most Coral Sea reefs. Further,
the few exploratory studies of MCEs in the CSMP are
restricted to a select number of reefs and have all focused
on benthic organisms, specifically scleractinian corals.
To date, there have been no quantitative surveys of fish
communities in the CSMP, or the Coral Sea more broadly,
at depths below 20 m. The ecology and biodiversity of
mesophotic coral reef fish communities in the Coral Sea
are therefore scarcely known.
Two major global marine biogeographical regions meet
in the Coral Sea; the Southwest Pacific and the Central
Indo-Pacific, the latter of which includes the Coral Tri-
angle (Kulbicki etal. 2013) (Fig.1b). There has been
increasing recognition that “peripheral habitats” (typi-
cally isolated archipelagos, marginal seas and seamounts)
in regions surrounding major biodiversity centres can also
export biodiversity and connect biogeographical regions,
rather than function only as isolated population sinks
(Bowen etal. 2013; Simon etal. 2022). Over the past
10–20 years, updated species checklists from locations in
the Southwest Pacific have listed occurrence records for
several Indo-Pacific fishes not previously reported from
these areas (Randall etal. 2003; Fricke etal. 2011b, a).
Additionally, aquarist collections from one location in
the central Coral Sea Marine Park (Holmes Reefs) have
also reported deep-water fishes typically known only from
either the Central Indo-Pacific or Central Pacific regions
(Fenton Walsh, pers. com). Large-scale connectivity pat-
terns are not well established in the Coral Sea, but these
observations suggest greater connectivity between the
Central Indo-Pacific and Western Pacific region through
the Coral Sea than may currently be known. This may
be particularly true for mesophotic species that are not
recorded in shallow reef community surveys but utilise
deeper reef habitats.
In this study, we used Remotely Operated Vehicles
(ROVs) and Baited Remote Underwater Video systems
(BRUVs) to survey fish and benthic communities in the
Coral Sea Marine Park at depths between 1 and 100 m.
Here, we present the first occurrence records of 16 spe-
cies from quantitative ecological surveys of coral reef
fishes from seamounts in the Coral Sea. We also compare
fish species richness between shallow-water monitoring
surveys and ROV/BRUV surveys and identify a range of
mesophotic habitats found in the CSMP during this study.
We discuss these observations in the context of regional
connectivity, biogeography and the importance of sea-
mounts in tropical coral reef seascapes.
Fig. 1 a Location of the Coral
Sea relative to major marine
biodiversity regions adapted
from dissimilarity analysis
between shallow species
occurrence data by Kulbicki
etal. (2013) and Spalding etal.
(2007) for nested ecoregions
within bioregions; b Bathym-
etry of the Coral Sea from the
same region bounded in red as
in a (data from Beaman 2012).
Most seamounts in the Coral
Sea Marine Park can be seen
in a chain on the Queensland
plateau, offshore to the east
from the Great Barrier Reef.
The Southwest Pacific region
in the Coral Sea extends east
to New Caledonia and north to
the border of the Coral Triangle
and Papua New Guinea. The
northern seamounts of the Coral
Sea extend to the border with
the Central Indo-Pacific region,
west through the Torres Strait.
Arrows indicate conceptual
borders and connectivity with
adjacent biogeographical
regions

Marine Biodiversity (2024) 54:17 17 Page 4 of 18
Material andmethods
Four separate voyages in the Coral Sea Marine Park were
undertaken in 2021, 2022 and 2023. ROVs and BRUVs
were used to survey fish communities and benthic habitats
at 17 reefs and between 1 and 100 m (Fig.2). Single-camera
BRUVs were deployed for 1 h following the standard oper-
ating procedures outlined in Langlois etal. (2020). ROVs
(BlueRobotics BlueROV2) were fitted with a forward-facing
stereo-video system (SVS) to enable length estimates to be
made. SVS cameras (Paralenz or GoPro Hero 8 systems)
were calibrated prior to surveys using the software CAL and
the associated calibration method (SeaGis Pty, Australia).
ROV transects, each 30 × 5 m, were conducted parallel to
the reef contour using a timed swim method (ROV speed
0.2 m/s for 2 min 30 s) at a constant depth (+/− 2 m). For
each ROV deployment, two transects were conducted within
each 10-m depth band, starting with the deepest transects
and working upwards to the shallows. Sufficient horizontal
and vertical separation was attained between transects and
between depth bands by the known speed and time of the
ROV. Three GoPro Hero8 cameras inside deep-rated T-hous-
ings were mounted facing outwards left and right and down-
wards on the ROV. These cameras were set to the timelapse
photo function (1 photo every 10 s), capturing an image of
the benthos every ~2 m (15 photos per transect). For fish
community data, videos were interrogated in EventMeas-
ure (SeaGIS, Pty Australia), and each individual fish enter-
ing the frame (BRUV) or transect field-of-view (ROV) was
identified to the lowest possible taxonomic resolution. For
ROV stereo-video footage, length estimates were also made
of individual fishes in each transect (fork length) using the
software EventMeasure Stereo (SeaGIS, Pty Australia). All
major habitats at each reef were surveyed (outer reef, lagoon,
back-reef and reef passes between the lagoons and leeward
outer reef), with BRUVs mostly conducted in lagoons and
inner reef areas due to the steep sides of many Coral Sea
seamount reefs.
Fig. 2 Map of the Coral Sea,
East Coast Australia and survey
sites from this study. All sites in
the CSMP surveyed by BRUV,
ROV and shallow diver surveys
are labelled by text. Locations
with yellow stars indicate reefs
with new locality observa-
tions for coral reef fishes. The
Holmes Reefs, where several
of these species have been
previously noted by aquarium
fish collectors, are marked by a
red circle. Reefs marked with a
black circle indicate locations
where no new observations of
reef fishes were recorded.

Marine Biodiversity (2024) 54:17 Page 5 of 18 17
We compared species records from shallow underwater
visual census monitoring surveys conducted by divers on
SCUBA (1–10 m depth) at the same reefs during the same
voyages (Hoey etal. 2021, 2022; Galbraith etal. 2022), as well
as fish species records from previous surveys of the same reefs
conducted by the Reef Life Survey Foundation (Edgar and Stu-
art-Smith 2014). Occurrence records and locations for species
only recorded by BRUV and ROV surveys in this study were
obtained from the Ocean Biodiversity Information System
(OBIS 2023a) and Global Biodiversity Information Facility
(GBIF.org 2023a) and were cross-referenced with other online
databases; Eschmeyer’s Catalogue of Fishes (Fricke etal.
2023), Atlas of Living Australia (ALA 2023), Fishes of Aus-
tralia (Bray and Gomon 2023), Reef Life Survey (RLS 2023),
FishBase (Froese and Pauly 2023), CSIRO Codes for Austral-
ian Aquatic Biota (Rees etal. 2023) and the Australian Faunal
Directory (ABRS 2020) as well as taxonomic experts. Known
depth records for all fishes recorded were also extracted from
FishBase using the rfishbase package (Boettiger etal. 2012)
and compared to depths at which they were observed by ROV
and BRUV surveys. To illustrate notable range extensions for
three species, previous extant range extents were calculated as
Extent of Occurrence (EOO) based on records obtained from
the aforementioned databases and plotted in R (R Core Devel-
opment Team 2023) using the packages ggmaps (Kahle and
Wickham 2013) and maps (Becker etal. 2022).
Results
A total of 274 ROV transects and 108 BRUV drops
were analysed and cumulatively recorded 361 species of
fishes from 41 families from depths between 1 and 100
m. Of these 361 species, 73 were recorded exclusively
by BRUVs, 105 exclusively by ROV and the remaining
183 were recorded by both methods (Online Resource 1).
Of the total 361 fish species, 128 (36%) were observed
at depths below their reported maximum known depth
as listed on the FishBase database (Online Resource
2). Thirty-four of these depth record extensions are for
species observed at depths greater than double their
previously reported maximum depth.
Compared to available data from shallow (< 10
m) underwater visual census surveys in the Coral
Sea (Ceccarelli etal. 2013; Stuart-Smith etal. 2013;
Hoey etal. 2020,2021,2022), this study recorded 50
additional species. Prior to this study, thirteen of these
50 species were previously only known from a single
location in the central Coral Sea through observations
and/or collections by aquarium fish collectors at the
Holmes Reefs (F.Walsh pers. Com). Outside of these
collections, these are the first observations of these
thirteen species from quantitative fish community
surveys in the Coral Sea Marine Park, and broader
Coral Sea region (Table1).
A further three species have single records from
Boot (Anampses melanurus, Atlas of Living Australia,
2023a), Frederick (Mulloidichthys pfluegeri, Atlas of
Living Australia, 2023b) and Osprey (Valenciennea
helsdingenii, Australian Faunal Directory 2023) reefs
respectively, but were recorded in our surveys at 11
other reefs (Table1). These records increase the range
extent of these three species in the region by between 4
and 12° of latitude.
Four species recorded by ROV in the CSMP represent
notable range extensions based on previous global occur-
rence records. Hoplolatilus randalli (Allen, Erdman &
Hamilton,2010), a relatively newly described species of
tile fish (family Malacanthidae), is currently known only
from Indonesia, the Philippines, Palau, Yap and the Solo-
mon Islands (Froese and Pauly 2023). A total of eight indi-
viduals were recorded at reefs spanning the northern and
central CSMP (Ashmore and Lihou Reefs and East Dia-
mond Islet), all at depths below 70 m (Table1, Fig.3a). We
mostly observed H. randalli in pairs beside large mounds
of rubble, apparently built by the fish over their burrows.
The observations from this study are the southernmost
occurrence records for the species and expand the known
extent of occurrence for H. randalli by almost 10° of lati-
tude. Cephalopholis polleni (Bleeker,1868) was previously
only known in Australian waters from the Cocos (Keeling)
and Christmas Islands in the Indian Ocean. Elsewhere, C.
polleni, (family Serranidae), occurs at scattered localities
on oceanic islands across the Indian Ocean and wider Indo-
Pacific (Bray 2023). In the Coral Sea, this study recorded
one individual C. polleni at 97 m under a ledge at Osprey
Reef by ROV survey (Fig.3b), and it has also been col-
lected at Holmes Reefs (Fenton Walsh pers. com). These
Coral Sea records extend the southern range of C. polleni
in the Southwest Pacific by 6° of latitude. Pseudanthias
flavicauda (Randall & Pyle,2001) was recorded by ROV
survey at Osprey and Bougainville reefs in the northern
Coral Sea. P. flavicauda (family Serranidae) is known
from the Central and Southwest Pacific (Bray 2022; Fro-
ese and Pauly 2023), and recently from Tonga (Fricke etal
2011b) and New Caledonia (Fricke and Kulbicki 2007).
We observed abundant schools of P. flavicauda between 80
and 100 m at Osprey and Bougainville Reefs, and although
also collected from the Holmes Reefs (Fenton Walsh pers.
com), the observations from this study are the most west-
ern records for this species and the most northern extent
in the Coral Sea (Fig.3c). Bodianus paraleucosticticus
(Gomon,2006) was found in ROV surveys at Lihou and
Osprey Reefs at depths between 70 and 90 m. Together
with collections from Holmes Reefs, these new observa-
tions of B. paraleucosticticus (family Labridae) extend the

Marine Biodiversity (2024) 54:17 17 Page 6 of 18
Table 1 New occurrence records of 16 fishes from mesophotic depths on seamount reefs in the Coral Sea. Previous range extents and distributions are based on georeferenced occurrence data
from online databases (see methods) and personal communication with taxonomic experts and aquarium trade collectors. Reported depth ranges were obtained from FishBase, and depth of
observations from this study was presented for comparison
Species Family Known range New Coral Sea records Reported
depth range
(FishBase)
Observed
depth (this
study)
Hoplolatilus randalli 1 (Allen, Erdman &
Hamilton, 2010)
Malacanthidae Indo-West Pacific: Indonesia, Philippines,
Palau, Yap Islands and the Solomon Islands.
Lihou, East Diamond, Boot, Ashmore Reefs 30–85 m 70–85 m
Cephalopholis polleni 1 (Bleeker, 1868) Serranidae Indian Ocean and Western Pacific: Comoros
Islands to the Line Islands and French
Polynesia. It has not been recorded from
continental Australia or the major islands of
Indonesia. In Australia, it has been recorded
from Christmas Island and Cocos (Keeling)
Island.
Osprey Reef 30–120 m 91 m
Pseudanthias flavicauda 1, (Randall & Pyle,
2001)
Serranidae Western Central Pacific: New Caledonia, Fiji,
Tonga, Vanuatu and Tahiti.
Osprey, Lihou and Bougainville Reefs 30–61 m 92 m
Liopropoma sp. “Yellow tail” 1Serranidae Currently undescribed but known to have a
wide distribution in the Western Pacific
(Tea, pers com).
Osprey Reef Na 70–94 m
Bodianus paraleucosticticus 1 (Gomon, 2006) Labridae Western Pacific: Cook Islands and Papua
New Guinea; New Caledonia.
Osprey and Lihou Reefs 25–115 m 70–90 m
Hoplolatilus marcosi 1 (Burgess, 1978) Malacanthidae Western Pacific Ocean; Indonesia,
Philippines, Palau, Papua New Guinea and
the Solomon Islands. One report from Great
Barrier Reef shelf break with Coral Sea
(Sih etal. 2017).
Lihou, East Diamond, Ashmore Reefs 18–80 m 60–85 m
Abalistes filamentosus 1 (Matsuura &
Yoshino, 2004)
Balistidae Indo-West Pacific: Southern Japan to northern
and north-western Australia and New
Caledonia.
Chilcott Reef 60–180 m 65–85 m
Pogonoperca punctata 1 (Valenciennes, 1830) Serranidae Indo-Pacific: Comoros to the Line,
Marquesan and Society islands, north to
southern Japan, south to New Caledonia.
Indian Ocean: southern Natal, South Africa
and the Australian territories of Christmas
Island and Cocos (Keeling) Islands. In
Australia from South of Evans Shoal,
Northern Territory.
East Diamond Islet 10–216 m 53 m
Valenciennea helsdingenii 2 (Bleeker, 1858) Gobiidae Indo-West Pacific: southern Red Sea and
East Africa to Indonesia, north to southern
Japan, south to the Great Barrier Reef.
Solomon Islands. Known from Osprey Reef
in the Coral Sea.
East Diamond Islet 1–45 m 52 m

Marine Biodiversity (2024) 54:17 Page 7 of 18 17
Table 1 (continued)
Species Family Known range New Coral Sea records Reported
depth range
(FishBase)
Observed
depth (this
study)
Xanthichthys auromarginatus 1 (Bennett,
1832)
Balistidae Ind-Pacific: From Mauritius east to northern
Australia to Hawaii and Society Islands.
North to Southern Japan. Northern Great
Barrier Reef (Escape Reef), in Christmas
and Cocos (Keeling) Islands and Lord
Howe Island.
Willis Island, Osprey, Bougainville, Mellish,
Ashmore, Boot and Wreck Reefs
8–140 m 40–95 m
Anampses melanurus 3 (Bleeker, 1857) Labridae Pacific Ocean: Indonesia to the Marquesas
and Society Islands, north to Ryukyu
Islands, south to Scott Reef Western
Australia. Range extends to Easter Island.
Ashmore, East Diamond, Osprey, Holmes,
Wreck Reefs
15–40 m 72 m
Pyronotanthias aurulentus 1(Randall &
McCosker, 1982)
Serranidae Eastern Central Pacific: Line Islands Osprey and Bougainville Reefs ?–52 m 92 m
Genicanthus bellus 1 (Randall, 1975) Pomacanthidae Western Pacific; Tahiti, Guam, Palau, Tonga,
the Cook Islands, the Marshall Islands, the
Philippines, southern Japan and southern
Indonesia. Indian Ocean of the Cocos
(Keeling) Islands and Christmas Island.
Osprey and Bougainville Reefs 25–100 m 66–82 m
Pycnochromis leucura 1 (Gilbert, 1905) Pomacentridae Indo-west-central Pacific, including
Madagascar, Mascarenes, Réunion,
Mauritius, Andamans, Indonesia, Hawaiian
Islands, Marquesas Islands and Gambier
Islands, southern Japan and Ryukyu Islands,
eastern Indonesia, New Caledonia.
Osprey Reef 20–120 m 84 m
Cirrhilabrus roseafascia 1 (Randall &
Lubbock, 1982)
Labridae Western Central Pacific: New Caledonia,
Philippines, Fiji, Palau, Samoa and Vanuatu.
Australia: Myrmidon region of the GBR (Sih
etal. 2017).
Osprey Reef, East Diamond Islet 30–155 m 94 m
Mulloidichthys pfluegeri 4 (Steindachner,
1900)
Mullidae Indo-West Pacific: Reunion, Hawaiian,
Marquesan and Society Islands. Eastern
Indonesia, north to Southern Japan and
south to Tonga. Known from the GBR in
Australia.
Chilcott, Flinders, Herald, Holmes, Kenn,
Lihou, Willis, Wreck Reefs
30–110 m 40–67 m
Superscript numbers next to species names indicate observations from three other single CSMP locations 1Holmes Reefs (F.Walsh Pers. com) 2Osprey Reef (Australian Faunal Directory 2023),
3Boot Reef (Atlas of Living Australia, 2023a, b), 4Frederick Reef (Atlas of Living Australia 2023a, b)

Marine Biodiversity (2024) 54:17 17 Page 8 of 18
previously reported distribution west from New Caledo-
nia and south from Papua New Guinea into the Coral Sea
(Fig.3d).
The presence of mesophotic coral ecosystems was con-
firmed at all 17 of the reefs surveyed by ROV and BRUV
(Fig.4). Several sites possessed remarkable hard coral cover
Fig. 3 Current extent of occurrence plotted as coloured hulls for
a Hoplolatilus randalli; b Cephalopholis polleni; c Pseudanthias
flavicauda; d Bodianus paraleucosticticus. Occurrence data were
obtained from OBIS (2023b,c,d,e) and GBIF (2023b,c,d,e). New
observations of each species from the Coral Sea by this study are rep-
resented by yellow stars

Marine Biodiversity (2024) 54:17 Page 9 of 18 17
(preliminary estimates ~70–85%) at depths between 50 and
100 m (Fig.5a and d). Multiple other non-coral dominated
mesophotic habitats were also found including Cycloclypeus
fields (large benthic foraminifera) (Fig.5b), octocoral domi-
nated walls (Fig.5c), seagrass (Fig.5e) and extensive Hal-
imeda meadows (Fig.5f).
Discussion
The new occurrence records presented here span a
considerable latitudinal gradient and provide evidence
of more widespread distributions for multiple fishes at
mesophotic depths in the region than previously known.
These observations are consistent with the role of seamounts
as stepping-stones for mesophotic fishes within the Coral
Sea and between other neighbouring biogeographic regions.
The provision and amount of suitable habitats at mesophotic
depths throughout the seamount chain, together with the
spatial arrangement of seamounts across the Coral Sea basin,
are likely key mechanisms supporting a stepping-stone
model of ecological connectivity in the region.
Total habitat area and the arrangement of a variety of
habitat types are fundamental components of species-area-
isolation relationships that drive biodiversity patterns (Mac-
Arthur and Wilson 1967; Connor and McCoy 1979; Fahrig
2013; Hanski 2015). The presence of multiple deep-water
habitats at individual Coral Sea reefs highlights that there is
considerably greater habitat area and habitat heterogeneity
within the Coral Sea than known from shallow reefs alone.
Fig. 4 Twelve species of coral reef fish recorded by ROV and BRUV
surveys in the Coral Sea Marine Park (CSMP) between depths of
50 and 100 m. All are previously known from one reef location in
the CSMP but are reported here from multiple other reefs spanning
the full latitudinal extent of the CSMP. a Pogonoperca punctata; b
Pycnochromis leucura; c Valenciennea helsdingenii; d Abalistes fila-
mentosus; e Liopropoma sp. “yellow tail”; f Xanthichthys auromar-
ginatus; g Mulloidichthys pfluegeri; h Anampses melanurus; i Cir-
rhilabrus roseafascia; j Genicanthus bellus; k Hoplolatilus marcosi;
l Pyronotanthias aurulentus. C.roseafascia, M.pfluegeri and H.mar-
cosi have been recorded on the outer GBR shelf break (Sih etal.
2017) but were previously not confirmed to be widely distributed
throughout the Coral Sea

Marine Biodiversity (2024) 54:17 17 Page 10 of 18
This is significant given that most metrics of ecological iso-
lation comprise some measure of patch size combined with
distance from nearest neighbouring habitat and the prop-
erties of the surrounding matrix (Moilanen and Nieminen
2002; Prugh etal. 2008). Compared to continental scales
(thousands of kilometres), seamounts in the Coral Sea are
separated by relatively small distances (< 450 km maxi-
mum distance). Isolation can both positively and negatively
affect biodiversity, either via demographic effects (Hanski
etal. 2013; Fahrig 2013; Jones etal. 2020) or distance from
anthropogenic influences (Demartini etal. 2008; Williams
etal. 2011; Bennett etal. 2018). In the context of this study,
the relatively small distance between many reefs throughout
the seamount chain, in combination with increased habitat
area and heterogeneity at mesophotic depths, may represent
an optimal level of isolation between populations. This in
turn would facilitate dispersal and enhanced connectivity
for some taxa through the stepping-stone model (Baum etal.
2004; Saura etal. 2014).
The relationships between increased habitat area
and reduced isolation, together with levels of habitat
heterogeneity and quality, also drive biodiversity through
other ecological dynamics (Gratwicke and Speight 2005;
Szangolies et al. 2022). For example, Halimeda spp.
meadows, seagrass and other macroalgae habitats are known
to provide valuable nursery habitats for reef fish settlement
and recruitment (Sambrook etal. 2019; Tang etal. 2020;
Sievers etal. 2020). Although this function has not been
extensively tested in MCEs, Halimeda meadows are known
to support diverse mesophotic fish communities (Langston
and Spalding 2017; Spalding etal. 2019), and we found these
habitats on deep outer reef slopes and in lagoons of all the
Coral Sea reefs surveyed. Given their isolation from other
coastal nursery habitats, these habitats may be particularly
important for the early life stages of fishes and invertebrates,
and thereby the replenishment and maintenance of Coral
Sea populations. Similarly, areas of high coral cover at
mesophotic depths increase total habitat area and resource
availability for coral-associated and dependent fishes.
Although resources at range margins, including depth, can
be of lower quality and affect the physiological condition of
some reef fishes (Munday 2001; Srinivasan 2003; Hoey etal.
2007), others including highly specialised obligate coral-
feeding butterflyfishes have been shown to access equal or
greater resources from deeper reefs without impacting their
fitness (MacDonald etal. 2018; MacDonald etal. 2019).
Fig. 5 Varied mesophotic coral
ecosystems and other meso-
photic habitats in the Coral Sea
a areas of high coral cover, 77
m, Lihou Reef; b Oxychellinus
orientalis and Cirrhilabrus
bathyphilus on a dense slope
of Cycloclypeus at 82 m, East
Diamond Islet; c Oxycheilinus
orientalis on a steep wall with
soft corals and gorgonians,
94 m, Osprey Reef; d high
abundance of fishes with high
and complex coral cover, 57 m
Bougainville Reef; e seagrass
(Halophila gradients) at 43 m
East Diamond Islet; f dense
Halimeda meadows at 67 m,
Lihou Reef

Marine Biodiversity (2024) 54:17 Page 11 of 18 17
Further, other studies of energetic trade-offs associated
with marginal coral reef habitats have found that deep reefs
support robust subpopulations through demographic and
reproductive plasticity (Goldstein etal. 2016). Biological
traits including habitat preference, diet and dispersal ability
therefore also strongly influence connectivity differentially
between species and ontogenetic stages (Hixon and Jones
2005; Goldstein etal. 2017). In the context of this study,
this is evident from the function of seamounts as stepping-
stones for deep-sea invertebrates, where mismatches
between expected and observed dispersal patterns can be
explained by a combination of environmental parameters
and biological traits (Miller and Gunasekera 2017).
The occurrence of several fishes in the Coral Sea which
were previously only known from either the Indo-Pacific
region or South and Western Central Pacific also aligns with
other ecological hypotheses explaining larger-scale patterns
in marine biodiversity. “The biodiversity feedback” theory
proposes that peripheral regions actively contribute to the
export of taxa and lineages back to biodiversity hotspots,
rather than acting only as sink populations (Bowen etal.
2013). Range expansions into the Coral Sea from both the
Central Indo-Pacific (e.g. H. randalli) and the Western
Central Pacific (e.g. P. flavicauda) suggest multiple
directions of connectivity between these biogeographical
regions through the Coral Sea. The proximity of the Coral
Sea to the global centre of marine biodiversity, the Coral
Triangle in the Indo-Pacific, mean reef habitats in the Coral
Sea may represent a particularly important link in larger-
scale patterns of reef fish biodiversity, rather than isolated
populations with limited distribution (Hobbs etal. 2009;
Budd and Pandolfi 2010). Though often speculated, prior to
this study, there has been scant empirical evidence to support
the role of the Coral Sea seamounts as stepping-stones for
reef fish populations (but see Van Herwerden etal. 2009),
or in contributing to biodiversity feedback between regions.
The stepping-stone model of dispersal has been shown
to contribute to the biodiversity feedback process for
reef fish assemblages in other regional seamount chains
(Pinheiro etal. 2015, 2018; Mazzei etal. 2021). From
studies in the Southwest Atlantic, coastal populations
closer to the Brazilian continental shelf represent areas of
higher biodiversity, and genetic connectivity exists in both
directions between these populations and the most offshore
seamounts (Simon etal. 2022). Interestingly, of the 16 new
occurrence records found in this study from the Coral Sea
Marine Park, 13 are not known from the neighbouring Great
Barrier Reef. At least nine of these species are deep-water
specialists, typically only known from depths greater than
30 m and up to 120 m. We include H. marcosi in these
deep-water specialists which, prior to these multiple new
Coral Sea observations, is only reported in Australia from
a single individual at 100 m at the GBR shelf break (Sih
etal. 2017) and has not been recorded from the reefs of the
GBR itself. Although a lack of observations on the GBR for
these 13 species may reflect low sampling from mesophotic
depths, the shallow geomorphology of the GBR shelf
(30–50 m, Hopley 2006) likely restricts the establishment
of populations of the deep-water specialist, or mesophotic,
species reported here. Indeed, the occurrence of these
mesophotic species throughout the full latitudinal extent of
the CSMP, but not on the GBR, suggests that connectivity
for these species is greater latitudinally along the seamount
chain, where deep-water habitat is available between the
Coral Triangle and island chains of the Southwest Pacific.
Connectivity patterns in the Coral Sea remain unclear but
for shallow-water studies, both genetic analyses (Planes etal.
2001; Payet etal. 2022) and dispersal-driven connectivity
models (Treml etal. 2008) suggest that connectivity between
the Coral Sea and GBR is generally weak. The barriers to
connectivity between these regions have not been fully
established, but the spatial separation of these two regions
by deep open water and the lack of mesophotic habitat on
the GBR would certainly contribute to a dispersal barrier
for mesophotic species from the Coral Sea seamounts to
the shallow GBR shelf. Further studies utilising genetic
sampling of mesophotic fishes with extended ranges
throughout the CSMP and from neighbouring regions are
required to test these aspects of the biodiversity feedback
hypothesis. Regional and localised oceanographic processes
will also determine the nature and direction of population
connectivity and barriers to dispersal throughout deep reefs
of the region. For example, dispersal via large-scale ocean
currents may be the main mode of seamount colonisation for
some taxa (Leal and Bouchet 1991), but localised seamount-
generated flows may be more important for explaining the
distribution of others (Richer de Forges etal. 2000). Finally,
although there is limited connectivity between shallow and
mesophotic ecological assemblages in many regions for
some taxa (Morais and Santos 2018; Bongaerts and Smith
2019; Stefanoudis etal. 2019), the depth extensions for 158
species in the Coral Sea reported by this study suggest that
boundaries between shallow and deep assemblages may be
shifted deeper here than community breaks known from
other regions (Lesser etal. 2019). Although MCEs clearly
warrant conservation actions and scientific investigation
independent from shallow-water reefs (Bridge etal. 2013;
Rocha etal. 2018), it is recognised that the degree of MCE
species overlap with shallow-water assemblages can vary
considerably by taxon and location (Laverick etal. 2018).
This necessitates further ecological and environmental
sampling from MCEs in remote geographic regions, like
the Coral Sea, and in understudied habitats, like seamounts.
A second mechanism, particularly relevant to patterns
of mesophotic fish diversity, is the “Habitat Persistence
Hypothesis” (HPH) (Copus etal. 2022). This theory posits

Marine Biodiversity (2024) 54:17 17 Page 12 of 18
that during periods of lower relative sea level, deeper marine
habitats can persist, particularly on complex bathymetries
where there are horizontal and low aspect areas, while
shallow-water habitats are dried out and communities here
are lost. Multiple biogeographical hypotheses of global pat-
terns and processes in reef fish diversity are supported to
varying degrees by often overlapping empirical evidence
(Mora etal. 2003; Gaither and Rocha 2013; Bowen etal.
2013; Cowman etal. 2017). Yet, most of this evidence is
derived from shallow-water coral reefs (< 30 m) which are
estimated to represent only 20% of global coral reef habitat
(Pyle and Copus 2019). These data therefore must be con-
sidered incomplete regarding both species inventories and
the extent of available habitat. Indeed, unlike shallow-water
reef fishes, diversity for mesophotic fishes does not appear to
attenuate with distance from the Coral Triangle (Pyle 2000,
2005; Pyle and Copus 2019), and this mismatch suggests
that there are further mechanisms shaping reef fish diversity
than have currently been considered (Pinheiro etal. 2023).
Again, although the lack of mesophotic sampling effort in
the region must be acknowledged, the absence of fishes
which are reported in this study from the adjacent GBR
aligns with several mechanisms proposed by the HPH. Dur-
ing periods of lower relative sea level, species richness may
have been retained in deeper habitats of the Coral Sea, and
in these persisting habitats, evolutionary processes would
continue among populations, potentially driving higher rates
of speciation. Certainly, taxonomic revisions and redefined
species complexes within some reef fish genera demonstrate
high levels of speciation across broad distributions which
span island chains, oceanic islands and seamounts in the
Western and Indo-Pacific. These include the Pomacentrus
philippinus group, which in the Coral Sea is represented
by P. imitator but in the adjacent GBR is P. magniseptus
(Allen etal. 2017), the goby genus Nemateleotris (Tea and
Larson 2023) and multiple Pseudanthias species (Anderson
2018, 2022; Gill 2022) many of which are deep-water spe-
cialists. The compilation and comparison of updated species
inventories is required to test the applicability of the HPH
in the Coral Sea, as well as genetic sampling to establish
patterns of connectivity between mesophotic populations in
adjacent regions. The complex bathymetries of the Coral Sea
seamounts do, however, constitute a significant system that
aligns with many of the mechanisms proposed by the HPH
and is a promising region in which to test these concepts
further.
Increased mesophotic surveys will likely continue to
increase diversity records for the region and comprehensive
fish species checklists for the Coral Sea Marine Park,
and Coral Sea region more broadly, will undoubtedly
continue to expand on the observations presented by this
study. Recent large-scale survey efforts in the region have
collected the most detailed bathymetry data for these reefs
to date (Carroll etal. 2021; Beaman etal. 2022; Brooke
and Schmitt Ocean Institute 2022) and have substantially
expanded our understanding of deep-sea habitats of the
Coral Sea. Unfortunately, despite the evident value of
deep-sea exploration, technical constraints on large ship-
based ROVs mean that such work often only focuses on
mesophotic habitats for short periods of time. Our findings
highlight the utility of small, affordable ROVs as an effective
tool for conducting mesophotic surveys in remote regions
where technical diving is often not feasible. Although
beyond the scope of this study, trait-based analysis of the
fish community data collected by ROV surveys would be
an informative line of further investigation to establish
how ecological characteristics (e.g. body size, depth range,
habitat preference, dispersal ability) contribute to habitat use
and geographical ranges throughout the Coral Sea. Further,
despite significant advances in understanding connectivity
and recruitment patterns in tropical reef fishes (Jones etal.
1999; Mora etal. 2003; Mora 2004; Planes etal. 2009;
Jones 2015; Almany etal. 2017), ecological studies from
seamounts are typically focused on cold-water, true deep-
sea taxa (Rowden etal. 2005, 2010; Pitcher etal. 2007;
Clark etal. 2010a; Rogers 2018). Coral reef communities
from tropical seamounts, including those inhabiting MCEs,
are therefore underrepresented in an already understudied
global marine habitat but are known to support abundant
and diverse reef fish communities (Letessier etal. 2019;
Galbraith etal. 2021; Leitner etal. 2021). As biodiversity
hotspots and important patch habitats for connectivity,
seamounts should more widely be considered global
conservation priorities in coral reef seascapes (McCook
etal. 2009; Riva and Fahrig 2022; Thompson etal. 2023).
Increased mesophotic community surveys in the Coral
Sea region and in other tropical seamount chains will
contribute to baseline knowledge of reef fish community
structure and species distributions in these habitats. Further
observational studies examining species turnover between
seamounts should also be supported by genetic sampling to
establish the nature and direction of population connectivity
throughout tropical seamount chains and within adjacent
biogeographical regions.
The findings from this study support the stepping-stone
model of dispersal for seamounts in a tropical seascape.
The new records of 16 fishes recorded throughout the
Coral Sea Marine Park confirm that the geographic range
of many tropical reef fishes is more widespread than
currently reported. Sampling deficiencies in the region
and at mesophotic depths are clearly a significant reason
behind new species observations. Nevertheless, these
new records from the Coral Sea contribute to evidence
that tropical seamounts are important habitats for reef
fish dispersal and connectivity between the Indo-Pacific,
Coral Triangle and Western Pacific region. The discovery

Marine Biodiversity (2024) 54:17 Page 13 of 18 17
of high coral-cover mesophotic reefs, combined with other
diverse deep-water benthic communities, demonstrates
the potential of mesophotic habitats of the Coral Sea to
provide valuable corridors for the dispersal of coral reef
fishes. The spatial arrangement of the Coral Sea seamounts
along the boundary of major biogeographical regions
also suggests that these tropical seamounts function in
processes driving large-scale marine biodiversity patterns.
These include but are likely not limited to biodiversity
feedback between peripheral regions and centres of
marine biodiversity and differences in the distribution and
diversity of mesophotic fishes driven by habitat persistence
through periods of sea-level change. These paradigms may
operate across other regional tropical seamount chains
and suggests these habitats have an important role in the
maintenance and regulation of global biodiversity patterns
for coral reef taxa.
Supplementary Information The online version contains supplemen-
tary material available at https:// doi. org/ 10. 1007/ s12526- 024- 01404-0.
Acknowledgements We are grateful to JH Choat, GP Jones and C
MacDonald for thoughtful discussions and comments on this manu-
script. Thanks also to D Bray, A Hay, M McNeil, J Pogonoski, YK Tea
and F Walsh for assistance with cross referencing species distributions
and identification. ROV and BRUV surveys were logistically supported
by the skippers of the MV Iron Joy, Rob Benn, and MV Argo, Jeff
Farnham, and their crew. Photo of B. paraleucosticticus in Fig.3d cour-
tesy of Richard Bajol/Alain Daoulas. We also thank three anonymous
reviewers for their engagement with and constructive feedback on our
manuscript. Field studies in the Coral Sea Marine Park were carried out
with permission from the Director of National Parks (permit number
PA2020-00092-3).
Funding Open Access funding enabled and organized by CAUL and
its Member Institutions Funding for all surveys was provided by the
Australian Government’s Our Marine Parks Grants (Rounds 2 and 3)
to ASH, GFG and ECM (grant numbers 4-FISKTNX and 4-HAY3RAP
respectively) and the ARC Centre of Excellence for Coral Reef Stud-
ies (ASH). Additional ROV surveys in July 2021 were facilitated by
Parks Australia with operational support from Queensland Department
of Environment and Science as part of the Coral Sea Island Health
voyage.
Declarations
Conflict of interest The authors declare no competing interests.
Ethical approval This research was conducted under James Cook Uni-
versity Animal Ethics permit A2721.
Sampling and field studies All necessary permits for sampling and
observational field studies have been obtained by the authors from the
competent authorities and are mentioned in the acknowledgements, if
applicable. The study is compliant with CBD and Nagoya protocols.
Data availability All species recorded by ROV and BRUV in this study
are provided as Online Resource 1. A list of depth extensions for fishes
compared to FishBase records is provided in Online Resource 1. Occur-
rence data used to plot Fig.3 were downloaded from the OBIS database
(OBIS 2023a,b,c,d,e), GBIF database (GBIF 2023a,b,c,d,e) and are
provided as a csv file at 10.5281/zenodo.7844209.
Author contribution All authors contributed to the study conception
and design. Material preparation and data collection were performed
GFG and BJC. Analysis was performed by GFG, BJC and ECM. The
first draft of the manuscript was written by GFG, and all authors com-
mented on previous versions of the manuscript. All authors read and
approved the final manuscript.
Open Access This article is licensed under a Creative Commons Attri-
bution 4.0 International License, which permits use, sharing, adapta-
tion, distribution and reproduction in any medium or format, as long
as you give appropriate credit to the original author(s) and the source,
provide a link to the Creative Commons licence, and indicate if changes
were made. The images or other third party material in this article are
included in the article’s Creative Commons licence, unless indicated
otherwise in a credit line to the material. If material is not included in
the article’s Creative Commons licence and your intended use is not
permitted by statutory regulation or exceeds the permitted use, you will
need to obtain permission directly from the copyright holder. To view a
copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
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