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Tropical seamounts as stepping-stones for coral
reef shes: range extensions and new regional
distributions from mesophotic ecosystems in the
Coral Sea, Australia
Gemma F Galbraith ( gemma.galbraith@jcu.edu.au )
James Cook University https://orcid.org/0000-0001-9888-2918
Benjamin J Cresswell
James Cook University https://orcid.org/0000-0003-0191-8828
Eva C McClure
James Cook University https://orcid.org/0000-0002-3073-1264
Andrew S Hoey
James Cook University https://orcid.org/0000-0002-4261-5594
Research Article
Keywords: seamounts, coral reef shes, Coral Sea, mesophotic coral ecosystems, biodiversity, ROVs,
BRUVs
Posted Date: September 15th, 2023
DOI: https://doi.org/10.21203/rs.3.rs-3342963/v1
License: This work is licensed under a Creative Commons Attribution 4.0 International License.
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Additional Declarations:
The authors have no competing interests to declare that are relevant to the content of this article
Tropical seamounts as stepping-stones for coral reef fishes: range
extensions and new regional distributions from mesophotic
ecosystems in the Coral Sea, Australia
Galbraith, G.F.*1,2 Cresswell, B.C.1,2, McClure, E.C.1,2 and Hoey, A.S.1,2
*Corresponding author gemma.galbraith@jcu.edu.au
1Marine Biology and Aquaculture, College of Science and Engineering, and
2ARC Centre of Excellence for Coral Reef Studies, James Cook University, Townsville, Queensland 4811,
Australia
Author ORCHID ID: G.F.G 0000-0001-9888-2918; B.J.C 0000-0003-0191-8828; E.C.M
0000-0002-3073-1264; A.S.H 0000-0002-4261-5594
Key words: seamounts, coral reef fishes, Coral Sea, mesophotic coral ecosystems,
biodiversity, ROVs, BRUVs
2
Abstract
Seamounts and remote oceanic islands provide 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 – 100m 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 sixteen 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 the 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 seventy-eight
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 thirteen 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
3
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.
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 3000m) (Davies et al. 1989; Bridge et al. 2019).
Ecologically, seamounts have been variously considered as either stepping stones for
dispersal that can promote regional connectivity (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).
Generalities for these contrasting hypotheses remain 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). Despite wide recognition as important marine habitats, in both tropical
and temperate oceans, seamounts remain one of the least explored and studied marine biomes
on earth (Clark et al. 2010b; Yesson et al. 2011) . Consequently, many regional seamount
chains require considerably greater sampling effort to establish patterns of biodiversity and
4
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 managed 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 combined protected area in the world (Director of National Parks 2018). In
Australia’s CSMP, over 30 individual reef systems are spread across 22 degrees of latitude on
the Queensland and Marion Plateaux and constitute ~24,000km2 of shallow water (< 30m)
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 (Figure 1). It
is therefore not surprising that the CSMP supports a high diversity of reef fish (>600 species)
and high abundance and biomass of sharks and other large predatory fishes (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 support 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 (Ceccarelli 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 likely a key component of ecological connectivity in this region and
with the wider Central Pacific Ocean.
5
Fig.1 a) Location of the Coral Sea relative to major marine biodiversity regions adapted from
dissimilarity analysis between species occurrence data by Kulbicki et al. (2013) and b)
Bathymetry of the Coral Sea from the same region bounded in red as in Figure 1a (data from
Beaman 2010). 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 boarder of the Coral
Triangle and Papua New Guinea. The northern seamounts of the Coral Sea extend to the
boarder with the Central Indo-Pacific region, west through the Torres Strait. Arrows indicate
conceptual borders and connectivity with adjacent biogeographical regions.
6
Although recent large-scale monitoring efforts (e.g. Hoey et al. 2020, 2022) and baseline
surveys (e.g. Ayling and Ayling 1985; Oxley et al. 2004; Edgar et al 2015) 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, restrictive
institutional and occupational diving regulations in Australia have led to a paucity of coral
reef research conducted in depths below 20m (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 2010). 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 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 benthic habitats between 0 -150m 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
7
of fish communities in the CSMP, or the Coral Sea more broadly, at depths below 20m. The
ecology and biodiversity of mesophotic coral reef fish communities in the Coral Sea is
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 Triangle (Kulbicki
et al. 2013) (Figure 1b). There has been increasing recognition that “peripheral habitats”,
typically 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 patterns are not well
established in the Coral Sea but these observations suggest greater latitudinal 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 which are not
recorded in shallow water reef 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 – 100m. Here we present the first occurrence records of sixteen
species from quantitative ecological surveys of coral reef fishes from seamounts in the Coral
8
Sea. We also compare fish species richness between shallow water monitoring surveys and
ROV/BRUV surveys and present 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 seamounts in tropical coral reef seascapes.
Methods
Four separate voyages in the Coral Sea Marine Park were undertaken in 2021, 2022 and
2023. Remotely Operated Vehicles (ROVs) and Baited Remote Underwater Video systems
(BRUVs) were used to survey fish communities and benthic habitats at 17 reefs and between
1 – 100m (Figure 2). Single camera BRUVS were deployed for 1 hour following the standard
operating 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 associated calibration method (SeaGis Pty Australia). ROV
transects, each 30 x 5m, were conducted parallel to the reef contour using a timed swim
method (ROV speed 0.2m/s for 2 minutes 30 seconds) at a constant depth (+/- 2m). For each
ROV deployment, two transects were conducted within each 10m 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 known speed and time
of the ROV. Three GoPro Hero8 cameras inside deep-rated T-housings were mounted facing
outwards left and right and downwards on the ROV. These cameras were set to the timelapse
photo function (1 photo every 10 seconds), capturing an image of the benthos every ~2m (15
photos per transect). For fish community data, videos were interrogated in EventMeasure
(SeaGIS, Pty Australia) and each individual fish entering the frame (BRUV) or transect field-
9
of-view (ROV) 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.
We compared species records from shallow underwater visual census monitoring surveys
conducted by divers on SCUBA (1 - 10m 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 Stuart-
Smith 2014). Occurrence records and locations for species only recorded by BRUV and ROV
surveys 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; Atlas of living Australia (ALA 2023), Fishes of Australia (Bray and
Gomon 2023), Reef Life Survey (RLS 2023), fishbase (Froese and Pauly, 2023), CSIRO
Codes for Australian 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 Development Team 2023) using the packages ggmaps (Kahle and
Wickham 2013) and maps (Becker et al. 2022).
10
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 observations 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.
11
Results
A total of 274 ROV transects and 108 BRUV drops were analysed and cumulatively recorded
360 species of fishes from 41 families (Online Resource 1). Compared to available data from
shallow (<10m) underwater visual census surveys, 50 additional species were recorded by
ROV and BRUV at depths between 1 – 100m. 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 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 2023), Frederick (Mulloidichthys pfluegeri, Atlas of Living Australia 2023) and
Osprey (Valenciennea helsdingeni, Australian Faunal Directory 2023) reefs respectively but
were recorded in our surveys at 11 other reefs (Table 1). These records increase their range
extent in the region by between 4 -12 degrees of latitude. Of the 360 fish species recorded by
BRUV and ROV surveys in this study, 158 (44%) were observed at depths below their
reported maximum known depth as listed on the fishbase database (Online Resource 2). 34 of
these depth record extensions are for species observed at depths greater than double their
previously reported maximum depth.
Table 1 New occurrence records of 16 fishes from mesophotic depths on seamount reefs in the Coral Sea. Previous range extents and 1
distributions are based on georeferenced occurrence data from online databases (see methods) and personal communication with taxonomic 2
experts and aquarium trade collectors. Reported depth ranges were obtained from fishbase and depth of observations from this study presented 3
for comparison. 4
Species
Family
Known range
New Coral Sea
records
Reported
depth range
(fishbase)
Observed
depth (This
study)
Hoplolatilus randali 1
(Allen, Erdmann &
Hamilton, 2010)
Malacanthidae
Indo West Pacific: Indonesia, Philippines, Palau,
Yap Is. and the Solomon Islands.
Lihou, East
Diamond, Boot,
Ashmore Reefs
30 -85m
70m – 85m
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 -120m
91m
Pseudanthias flavicauda 1,
(Randall and Pyle 2001)
Serranidae
Western-central Pacific: New Caledonia, Fiji,
Tonga, Vanuatu and Tahiti.
Osprey, Lihou
and
Bougainville
Reefs
30 – 61m
92m
Liopropoma ‘sp. Yellow tail’ 1
Serranidae
Currently undescribed but known to have a wide
distribution in the Western-Pacific (Tea, pers
com).
Osprey Reef
Na
70-94m
Bodianus paraleucosticticus 1
(Gomon 2006)
Labridae
Western Pacific: Cook Islands and Papua New
Guinea; New Caledonia.
Osprey and
Lihou Reefs
25 -115m
70 – 90m
Hoplolatilus marcosi 1
Malacanthidae
Western Pacific Ocean; Indonesia, Philippines,
Lihou, East
18 – 80m
60 – 85m
13
(Burgess 1978)
Palau, Papua New Guinea and the Solomon
Islands. One report from Great Barrier Reef shelf
break with Coral Sea (Sih et al. 2017).
Diamond,
Ashmore Reefs
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 – 85m
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 – 216m
53m
Valenciennea helsdingeni 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 - 45m
52m
Xanthichthys auromarginatus 1
(Bennett 1832)
Balistidae
Indo 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 – 140m
40 – 95m
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 -40m
72m
Pyronotanthias cf aurulentus 1
(Randall and McCosker 1982)
Serranidae
Eastern Central Pacific: Line Islands
Osprey and
Bougainville
Reefs
? – 52m
92m
Genicanthus bellus 1
Pomacanthidae
Western pacific; Tahiti, Guam, Palau, Tonga,
Osprey and
25 – 100m
66 – 82m
14
(Randall 1975)
the Cook Islands, the Marshall Islands, the
Philippines, southern Japan and southern
Indonesia. Indian Ocean of the Cocos (Keeling)
Islands and Christmas Island.
Bougainville
Reefs
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 -120m
84m
Cirrhilabrus roseafascia 1
(Randall and 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 – 155m
94m
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 -110m
40 – 67m
Superscript numbers next to species names indicate observations from three other single CSMP locations 1Holmes Reefs (F.Walsh Pers.com)
5
2Osprey Reef (Australian Faunal Directory 2023), 3Boot Reef (Atlas of Living Australia 2023), 4Frederick Reef (Atlas of Living Australia 2023).
6
Four species recorded by ROV in the CSMP represent notable range extensions based on
previous global occurrence records. Hoplolatilus randalli (Allen et al. 2010), a relatively
newly described species of tile fish (family Malcanthidae), is currently known only from
Indonesia, the Philippines, Palau, Yap and the Solomon Islands (Froese and Pauly 2023). A
total of eight individuals were recorded at reefs spanning the northern and central CSMP
(Ashmore and Lihou Reefs and East Diamond Islet), all at depths below 70m (Table 1, Figure
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 degrees of latitude. 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 97m under a ledge at Osprey Reef by ROV survey
(Fig 3b) and it has also been collected at Holmes Reef (Fenton Walsh pers. com). These
Coral Sea records extend the southern range of C. polleni in the Southwest Pacific by 6
degrees of latitude. Pseudanthias flavicauda (Randall and 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 South West Pacific (Bray 2022; Froese and Pauly
2023), and recently from Tonga (Fricke et al 2011) and New Caledonia (Fricke and Kulbicki
2007). We observed abundant schools of P. flavicauda between 80 -100m 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 western records for this species and the
most northern extent in the Coral Sea (Figure 3c). Bodianus paraleucosticticus (Gomon
2006) was found in ROV surveys at Lihou and Osprey Reefs at depths between 70 – 90m.
16
Together with collections from Holmes Reefs, these new observations of B.
paraleucosticticus (family Labridae) extend the previously reported distribution west from
New Caledonia and south from Papua New Guinea into the Coral Sea.
Fig.3 Current extent of occurrence plotted as colored hulls for a) Hoplolatilus randalli and b)
Cephalopholis polleni c) Pseudanthias flavicauda and 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 represented by yellow stars.
17
In addition to range extensions, we found twelve other species, previously only known from a
single location in the Coral Sea, at multiple other reefs in ROV and BRUV surveys. None of
these species have previously been recorded in shallow water community surveys in the
Coral Sea Marine Park and most are deep-water specialists occurring below depths of 50m
(Figure 4). Three of these species have been recorded on the GBR but were previously not
confirmed to be widely distributed throughout the Coral Sea.
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 to 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 helsdingeni d) Abalistes filamentosus e) Liopropoma “sp. yellow tail” f)
Xanthichthys auromarginatus g) Mulloidichthys pfluegeri h) Anampses melanurus i)
Cirrhilabrus roseafascia j) Genicanthus bellus k) Hoplolatilus marcosi and l) Pyronotanthias
cf aurulentus.
18
The presence of mesophotic coral ecosystems was confirmed at all 17 of the reefs surveyed
by ROV and BRUV. Several sites possessed remarkable hard coral cover (preliminary
estimates ~70 -85%) at depths between 50 – 100m (Figure 5a and d). Multiple other non-
coral dominated mesophotic habitats were also found including Cycloclypeus beds (large
benthic foraminifera) (Figure 5b), octocoral dominated walls (Figure 5c), seagrass (Figure
5e) and extensive Halimeda meadows (Figure 5f).
Fig.5 Varied mesophotic coral ecosystems and other mesophotic habitats in the Coral Sea a)
areas of high coral cover, 77m, Lihou Reef b) Oxychellinus orientalis and Cirrhilabrus
bathyphilus on a dense bed of Cycloclypeus at 82m, East Diamond Islet c) Oxycheilinus
orientalis on a steep wall with soft corals and gorgonians, 94m, Osprey Reef d) High
abundance of fishes with high and complex coral cover, 57m Bougainville Reef e) seagrass
(Halophila gradients) at 43m East Diamond Islet and f) Dense Halimeda meadows at 67m,
Lihou Reef.
19
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 support the stepping stone model of
ecological connectivity for seamounts within the Coral Sea and between certain neighbouring
biogeographic regions. 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 only
acting as sink populations (Bowen et al. 2013). Range expansions into the Coral Sea from
both the Central Indo-Pacific (e.g. H. randalii) 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
20
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 sixteen new occurrence records found in
this study from the Coral Sea Marine Park, thirteen 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 30m and up to 120m. 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 100m 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 thirteen species may reflect low sampling from mesophotic depths, the
shallow geomorphology of the GBR shelf (30 - 50m, 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 is required to test
21
aspects of the biodiversity feedback hypothesis and quantify the nature and direction of meta-
population connectivity throughout deep reefs of the region.
A second mechanism, particularly relevant to patterns of mesophotic fish diversity, is “The
Habitat Persistence Hypothesis” (HPH) (Copus et al. 2022). This theory posits that during
periods of lower relative sea level, deeper 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
patterns 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 (<30m) which are estimated to represent only 20% of global coral reef habitat (Pyle and
Copus 2019) . These data therefore must be considered 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. During 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 amongst 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
22
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 specialists. The compilation
and comparison of updated species inventories is required to test the applicability of the
Habitat Persistence Hypothesis 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.
Total habitat area and the arrangement of a variety of habitat types are also fundamental
components of species-area-isolation relationships which drive biodiversity patterns
(MacArthur 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. This is significant given that most measures of ecological isolation
comprise some measure of patch size combined with distance from nearest neighbouring
habitat and the properties 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 (< 450km maximum 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 close proximity of many reefs throughout the seamount chain, in combination
23
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 beds 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 beds 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
Coral Sea reefs. 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 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). 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
24
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). 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.
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
25
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 as
conservation priorities in coral reef seascapes (McCook et al. 2009; Santos and Morato 2010;
Riva and Fahrig 2022). Increased mesophotic community surveys in the 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 should also
be supported by genetic sampling to establish the nature and direction of meta-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 sixteen fishes recorded throughout the Coral Sea
Marine Park confirm that the geographic range of many tropical reef fishes are more
widespread than currently reported. Sampling deficiencies in the region and at mesophotic
26
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 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 dispersal for 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.
Acknowledgements
We are grateful to JH Choat, GP Jones and C MacDonald for thoughtful discussion of and
comments on this manuscript. 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 courtesy of Richard Bajol/Alain Daoulas.
27
Compliance with Ethical Standards
The authors have no competing interests to declare that are relevant to the content of this
article. 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 commented on
previous versions of the manuscript. All authors read and approved the final manuscript.
Funding
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 Studies (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.
Ethics and permitting statement
This research was conducted under permission from the Director of National Parks (PA2020-
00092-3) and the James Cook University Animal Ethics Committee (approval A2721).
Data availability statement
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 2. Occurrence data used to plot Figure 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.
28
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