Content uploaded by Sonia J. Rowley
Author content
All content in this area was uploaded by Sonia J. Rowley on Jun 27, 2017
Content may be subject to copyright.
Content uploaded by Sonia J. Rowley
Author content
All content in this area was uploaded by Sonia J. Rowley on Jan 27, 2017
Content may be subject to copyright.
1!
Voyage Report: Rapid Assessment of the Geologist and Lōʻihi Seamounts
Sonia J. Rowley
University of Hawai‘i at Mānoa, USA
‘Sea of tranquility’
(Top: Pisces IV descending. Bottom Lōʻihi vents, L. Lamar. Quote: G. Stone 2016)
HURL-CI Project KOK1616
September 2016
2!
Project Title: Geologist and Lōʻihi Seamount Rapid Assessment
Project Code: KOK1616
Vessel: R/V Ka'imikai-o-Kanaloa (KoK)
Location: South Hawai‘i, Geologist and Lōʻihi Seamounts
Period: 5th – 9th September 2016
Voyage Personnel: Dr. Greg Stone (Principal Investigator, CI), Dr. M. Sanjayan (Senior Scientist, CI),
Dr. Sonia J. Rowley (Senior Scientist, UH), Dr. Lida Teneva (Science Manager, CI), Dr. Alan Eustace
(Observer, Google), Glenn Baum (BF), Terry Kerby (Submersible Pilot, HURL), Maximilian
Cremer (Submersible Pilot, HURL), Dr. John R. Smith (GIS Mapping, UH), Luis Lamar
(Videographer, WHOI), Michael Garland (Observer, Photographer, BF), Caleb Jones (Observer,
Journalist, AP), Kevin Connor (Media Manager, CI).
Expedition Summary This
This research project aimed to explore and describe the benthic biodiversity and geological history of
the little-studied Geologist seamounts, Cook and McCall, as well as revisit Hawai‘i’s youngest and most
active seamount, Lōʻihi. A total of six submersible dives were conducted. These dives yielded six bait
stations, seven geological samples, six microbial samples, and seventeen coral specimens including one
gorgonian coral species new to science. Benthic surveys were conducted on each dive revealing that
Cook and McCall harbour a wealth of hidden biodiversity characteristic of Vulnerable Marine
Ecosystems (VMEs). Bait station scavenging communities consisted of synaphobranchid eels, Smooth
Lanternsharks, a variety of shrimp, and most notably, a Pacific Sleeper shark in Pele’s Pit at Lōʻihi.
Finally, after >5 years the Pisces submersibles returned to Lōʻihi, Hawai'i’s youngest seamount, to
explore changes in geomorphology, and chemosynthetic microbial mat community composition.
Preliminary observations revealed a reduction in active venting, however, areas of low temperature
(~12-30ºC) hydrothermal ‘finger’ and ‘beehive’ chimney vents were present on the southeastern wall of
Pele’s Pit. Part of the southeastern wall vents were obscured which may have been due to a landslide
between 2014–2016. Exploration within the ‘Prime Fe-Mn Crust Zone’ (PCZ) is critical in this time of
unprecedented technological advancement, in order to protect deep-sea seamounts and associate
biological communities and biogeochemical interactions from the expansive economic interest in deep-
ocean mining of mineral deposits. This project has and could continue – if supported – to reveal valuable
research and conservation dividends for some of the oldest, and most isolated, and unprotected,
seamounts on earth.
Background
The Pacific Ocean harbours the greatest number of seamounts (submerged mountains) of any ocean. Of
those explored, the Geologist Seamounts have been relatively understudied, and are considerably older
than the adjacent Hawaiian Island chain due to a difference in origin (Dymond & Windom 1968).
Previous evidence suggests that the Geologist seamounts date back to the late Cretaceous (~80 Ma;
Sager & Pringle 1987), were formed along the east Pacific Rise, and subsequently rafted northwest due
to the movement of the Pacific Plate. In contrast, the Hawaiian seamounts formed through a ‘hotspot’ in
the underlying mantle and not at the plate boundary. Lōʻihi, the youngest in the Hawaiian chain (~0.3
3!
Ma), is still active, with unique microbial mat communities (Karl et al. 1989, Donachie et al. 2004,
Emerson et al. 2017). This trip provides a unique opportunity to explore, in close proximity, some of the
oldest Central Pacific seamounts along with the youngest.
Seamounts rise over 1000 m from the seabed, altering the flow of ocean currents, and thus capturing
plankton and nutrients. This supports high densities of corals, sponges and fishes, classifying seamounts
as vulnerable marine ecosystems (VMEs). With almost 100 species of deep-sea coral known from the
Hawaiian Islands, and more likely awaiting discovery (Watling et al. 2011), exploration on these
isolated pockets of biodiversity is crucial. Increased current flow on ridged compared to conical
seamounts, also would suggest a higher density and biodiversity on McCall vs. Cook. Yet, the shallow
summits of adjacent seamounts, once characterised by rich benthic and shark communities, are
continually subject to long-line fisheries and disturbing quantities of abandoned equipment (T. Kerby
pers. comm.). In addition, between 800-2500 m depth cobalt-rich ferromanganese coat (Fe-Mn crusts)
the basalt, which increases in thickness with age, and is thus vulnerable to exploitation by the deep-sea
mining industry. Curiously, recent evidence suggests Fe-Mn crusts at the Geologist seamounts McCall,
Ellis, and Swordfish were surprisingly thin (<2-10 mm; Bell et al. 2015). Whether this pattern is
persistent across bathymetry, seamounts, and affects key benthic components is unknown.
Objectives
With seamounts such as Cook as yet unexplored, it is unclear what biodiversity may be at stake, and
perhaps even the true age of these seamounts. Therefore, the primary objective of this project was to
explore seamounts of contrasting age, origin, and potential fisheries impact, and within this to: (1)
survey and sample key benthic invertebrate taxa such as gorgonian corals for taxonomic, phylogenetic
and associate microbial community analyses, (2) identify and observe megafaunal-scavenging
communities through baited cameras, (3) collect rock samples to provide new insights into the
geological history of these seamounts, and (4) sample the distinct microbial mats at Lōʻihi for
extremophile culturing, and metagenomic and metatranscriptomic analyses.
Methods
Study Sites
The Geologist Seamounts are in fact guyots (flat-top seamounts, or tablemounts) that are situated >50
miles southwest of the Big Island of Hawai‘i (Figure 1). Lōʻihi Seamount lies 21 miles southeast of the
Big Island, and is the youngest and smallest volcano in the Hawaiian-Emperor chain. Dives were
conducted on two of the Geologist seamounts, Cook and McCall, as well as Lōʻihi using the Hawai‘i
Undersea Research Laboratory’s (HURL) manned submersibles Pisces IV (see opening figure) and V in
tandem from the support vessel, RV Ka‘imikai-o-Kanaloa. Each submersible dive was tracked using
TrackLink 5000HA USBL (LinkQuest Inc.) interfaced with HYPACK software for real-time display
and tracking operations.
Sample and Data Collection
Data were collected using onboard videography (onboard Insite Pacific MINI-ZEUS HDTV, and Pisces
V externally fitted custom built 4K Camera System, L. Lamar & E. Kovacs, AIVL) and observer
documentation. Video and observer surveys were conducted throughout the dive documenting fish and
mega-invertebrate benthic communities (e.g., gorgonian octocorals, scleractinian corals, sponges).
Taxonomic identification was based on key diagnostic morphological characters and, where appropriate,
collected and preserved for detailed taxonomic assessment. Coral specimens such as gorgonian corals
were collected, preserved (see Table 1) and stored at -80ºC for downstream analyses.
4!
Figure 1. Location map of submersible dives; the Geologist seamounts Cook and McCall ~100 miles
southwest of the Big Island of Hawai‘i, and Lōʻihi seamount 22 miles southeast of the Big Island.
Image modified from Eakins et al. (2003).
Megafaunal-scavenging communities were assessed through filmed bait stations. Typically two 1-gallon
bags, with wooden dowels for ease of opening with the submersible manipulators, containing 1-inch
chunks of adult skipjack tuna and squid were released onto the seafloor per station. Submersible lights
were switched off for ~5 minutes prior to observations. Megafaunal-scavenging communities were
quantified using criteria as outlined by Yeh & Drazen (2009); whereby active feeding or attraction to the
bait was identified as scavenging.
Chemosynthetic microbial mats characterising Lōʻihi were collected, as well as the adjacent water
column, and preserved for downstream analyses. Both microbial mats and water samples were also
plated on Eukaryote- targeted marine agar inoculated with antibiotics. Replicate plates were stored at 10
and 27ºC. Geological specimens (rocks) were collected, washed and dried at 60ºC for 48 hrs.
Data Analyses
Megafaunal-scavenging and ecological survey community data were analysed using multivariate
routines in R (R 2013). Morphological taxonomic analyses were conducted using an Olympus SZX16®
stereomicroscope and scanning electron microscopy (SEM) Hitachi S-800, for macro and micro-
morphological analyses respectively. Molecular analyses were conducted using methods as outlined by
Cook Seamount »
« Lōʻihi
«
McCall Seamount
5!
Table 1. KOK1616 expedition submersible dive summary and personnel.
Figure 2: Dive track lines for the Geologist seamounts (a) Cook, and (b) McCall. Red line 2000 m depth contour; ArcGIS Image: C. Kelley 2015.
Figure 2: Dive track lines for the Geologist seamounts (a) Cook, and (b) McCall. Red line 2000 m depth contour; ArcGIS Image: C. Kelley 2015.
Dive tracks inset by JR. Smith. Pisces IV – white track, Pisces V – blue track.
Date
Dive
Sub Personnel
Location
Max
Depth (m)
Bottom
Time
Dive
Duration
Pilot
Observer 1
Observer 2
Seamount
Lat
Long
06.09.2016
P4-298
M. Cremer
S. Rowley
M. Garland
Cook
N19:18.0
W157:10.0
1145
5.31
6.14
06.09.2016
P5-859
T. Kerby
G. Stone
C. Jones
Cook
N19:18.0
W157:10.0
1081
6.31
7.59
07.09.2016
P4-299
M. Cremer
S. Rowley
G. Stone
McCall
N18:42.9
W157:04.77
1053
4.24
5.51
07.09.2016
P5-860
T. Kerby
M. Sanjayan
A. Eustace
McCall
N18:42.9
W157:04.77
1156
6.29
7.51
08.09.2016
P4-300
M. Cremer
S. Rowley
M. Garland
Lōʻihi
N18:54.35
W157:15.70
1325
5.22
6.20
08.09.2016
P5-861
T. Kerby
G. Stone
A. Eustace
Lōʻihi
N18:54.35
W157:15.70
1325
6.22
8.39
Total (hrs)
34.39
42.54
6!
Figure 3: Lōʻihi Seamount in the Hawaiian Chain. ArcGIS Image: C. Kelley 2015. Dive tracks inset
by JR. Smith. Pisces IV – white track, Pisces V – blue track, P, Pele’s Pit.
McFadden et al. (2011, 2014). Metagenomic analyses on select gorgonian taxa, microbial mats and
adjacent water column were performed using Illumina MiSeq sequencing (see Yu et al. 2015; primers
from Leray & Knowlton 2016). Geological samples were thin sectioned and analysed as described by
Garcia et al. (2015). All specimens and subsequent data analyses are currently underway at the
University of Hawai‘i at Mānoa, USA.
Results
A total of six submersible dives over three dive days were successfully conducted during the 6-8th
September 2016, each up to eight hours duration culminating 43 hours dive time (Table 1, Figure 2 &
3).
Cook & McCall Seamounts Benthic Surveys
Benthic surveys (Figure 4, Appendix I) revealed unexpected clusters of biodiverse benthic
communities often characterising dramatic geological formations such as pinnacles, ledges and rift
walls. Such geological structures were frequently lined with diverse communities of suspension
feeding invertebrates, and more low relief terrain would often possess large patches of Calyptrophora
wyvillei Wright, 1885 colonies or numerous small (≤30 cm tall) colonies of Pleurocorallium
porcellanum (Pasternak, 1981). An increased abundance of sponges was present below 930 m depth
for both seamounts. Habitat forming (key stone) taxa such as gorgonians and sponges were host to a
7!
Figure 4. KOK1616 figure characterising the observed seamount environments and notable taxa. Cook
seamount: (a) striking ledges of invertebrate biodiversity and abundance, (b, c) two Grimpoteuthis cf.
‘dumbo’ octopus, (d) >3 m colony of Iridogorgia magnispiralis with Paramuricea n.sp., baring a
zoanthid, also Lepidisis, and Corallium in the background. McCall seamount: (e) separation along a
fracture encrusted in Fe-Mn, and bearing small colonies within the Coralliidae, (f) Hydrolagus cf.
purpurescens ‘Chimera’, (g) angular talus, (h) large Hemicorallium sp. on pinnacle edge. Lōʻihi
seamount (i) Pisces V descending into the ‘Pele’s Pit’ at marker 18, (j) marker 29, (k) sampling rocks at
marker 18 still with data loggers placed in 1997, (l) active beehive and finger vents (close up inset) on
the south east corner of Pele’s Pit, and (m) inactive vent (white). Images (e, g, i, j, m) M. Cremer, (a, d,
k) T. Kerby, (l & inset) M. Garland, (b, c, f, h) L. Lamar.
8!
variety of associate invertebrates particularly within the Ophiurida, Chirostylidae, and
Galatheidae. Biological communities, however, were not continuous, with areas of very
low abundance particularly where sediment was present. Substratum consisted of pillow and
columnar basalts, hard ferromanganese covered (Fe-Mn) rock, boulders, what appeared to be
unconsolidated talus, and small intermittent sediment pockets typically bearing current ripples.
Slopes were of high relief, and ancient dyke formations were often encountered. Loose talus was
not uncommon on the flanks of each seamount studied.
A thorough species inventory is currently underway (preliminary summary Appendix I),
however, conservative estimates consist of >100 species including >85 invertebrates (37 within the
Anthozoa) and 17 species of fishes for all seamounts surveyed. Overall numerous octocoral species
were present such as those within the precious coral family Coralliidae (e.g., P. porcellanum,
Hemicorallium) and Isididae (e.g., Lepidisis, Acanella), Acanthogorgia, and Paragorgia. A new
species of Paramuricea (Figure 6a) was discovered on both Cook and McCall, which aligns
closely, (but not completely), with the phenotypic traits of Paramuricea n.sp. B. in Muzik
(1979). Taxa within the Primnoidae (e.g., Calyptrophora, Candidella, Narella) as well as a
variety of Chrysogorgiidae; Chrysogorgia spp. including C. geniculata (Wright & Studer,
1889), Iridogorgia bella Nutting, 1908, and particularly large colonies of I. magnispiralis
Watling, 2007, I. splendens Watling, 2007, and Metallogorgia melanotrichos (Wright &
Studer 1889) were well represented throughout surveys conducted. Scleractinians such as
Enallopsammia, as well as unidentified zoanthids within the family Parazoanthidae were frequently
encountered particularly on dead or partially alive bamboo (Isididae) gorgonians. Numerous sponge
taxa were mainly within the Euretidae, Farreidae, and Euplectellidae. A variety of ascidians,
crinoids, hydroids (e.g., Stylasteriidae), Bathypathes (black coral), occasional Penatulids, and other
suspension feeding taxa were observed, along with Nereidaster bowersi Fisher, 1906 and other
members of the Echinodermata. Of the two sightings of the Dumbo octopus Grimpoteuthis
sp. at McCall, one was observed shedding its outer tissue layer, and it is unclear if both are of
the same genus. The scavenging fauna (e.g., Synaphobranchus, Etmopterus pusillus,
Nematocarcinus, Heterocarpus spp.) were also present in each survey. Other fish taxa
encountered included those within the Ophidiidae, Macrouridae (e.g., Coryphaenoides,
Nezumia), and the Chimera Hydrolagus cf. purpurescens (see Figure 4 for characteristic site and
taxon imagery).
Bait Stations In
In total six bait stations were conducted on each of the seamounts; Cook (n = 1: 127 mins duration,
23 individuals, 3 taxa, 11 individuals/hr), McCall (n = 3: 142 mins duration, 60 individuals, 5 taxa,
20 individuals/hr), and Lōʻihi (n = 2: 168 mins duration, 56 individuals, 5 taxa, 20 individuals/hr). A
formal observational analysis was not strictly possible due to obscured observations on some of the
dives; however, a thorough account of what was observed is provided below. Typical megafaunal-
scavenger taxa present (Figure 5, Appendix II) and actively feeding at all bait stations included eels
of the genus Synaphobranchus. The shrimp Nematocarcinus tenuirostris Spence & Bate, 1888, and
Heterocarpus spp. transported bait but were not observed to actively consume. Shrimp and juvenile
(≤25 cm in length) synaphobranchid eels were frequently observed attempting to consume and/or
transport chunks of bait notably larger than their anatomical capacity. Scavengers such as
Synaphobranchus, Heterocarpus, and Nematocarcinus, once in possession of bait, were observed to
swim at an upward angle, presumably to escape competition and/or predation. On two occasions
Synaphobranchus eels were observed snatching bait from the Heterocarpus shrimp, prior
to them swimming upwards at McCall. Of considerable note was an abundance of Smooth Lante-
9!
Figure 5. KOK1616 figure showing characteristic bait station fauna. Cook seamount: (a) Pisces V
deploying bait, (b) Synaphobranchus devouring bait. McCall seamount: (c) Pisces V deploying bait, (d)
Heterocarpus laevigatus carrying bait and Etmopterus pusillus. Lōʻihi seamount: (e)
Synaphobranchus carrying bait, (f, h) Somniosus pacificus inspecting Pisces IV and, (g) an
overambitious Heterocarpus sp. Images: (f, g) M. Garland, (a, c) T. Kerby, (b, d, e, h) L. Lamar.
10!
-rnsharks Etmopterus pusillus (Lowe, 1839) on the summits of Cook and McCall, yet barely
consumed any bait. The chimera Hydrolagus cf. purpurescens investigated the bait basket on Pisces
V at McCall but did not feed. Similarly in Pele’s Pitt, Lōʻihi, an adult Pacific sleeper shark
(Somniosus pacificus Bigelow & Schroeder, 1944) showed interest in both the submersibles and bait,
but did not feed.
Lōʻihi Seamount After
After more than five years this research trip enabled the Pisces submersibles to return to Lōʻihi and
successfully located Pele’s Pit (Figure 3), as well as numerous previously laid research location
markers (#18, 29, 36, 39, 48). These markers were typically coated with a ~1 cm thick layer of
orange-brown venting deposits (e.g., Figure 4i-k). Dike formations and sections of pillow lavas and
loose talus characterised the western upper flank of Pele’s Pit. All rock types were also covered in a
thin layer of orange-brown iron-rich venting deposits (Figure 4i-k, m, 6g). The bottom of Pele’s pit
(1324 m depth) was characterised by black volcanic sands and glass sediments bearing current ripples
and yellow/orange microbial flocks (Figure 5e-h). Microbial flocks originating from nearby active
vents settled in the depressions of the volcanic sediment ripple formations and crevices amongst
young talus. Active ‘finger’ chimney vents were observed (see cover page; Figure 4l, 6h-j), however,
none were as dramatic as those observed on former trips (Cremer M, Kerby T, pers. comm.).
Specifically, the previously active vents on the east wall of the pit at marker 36 at the North Hiolo
Ridge (see Glazer & Rouxel 2009) were only gently active. At marker 36 it was observed that a
landslide had likely covered the ‘Jet’ site from 2008 (Kerby T, pers. comm.). Active venting
increased from marker 39 at the Upper Hiolo Ridge southeastward ascending up the steep ridge wall
to and beyond marker 48, with an incredible forest of finger chimney vents. ‘Beehive’ vents were
also present (Figure 4l, 6i), both previously observed at Lōʻihi. There was a low incidence of
bacterial flocks and turbidity in the water column. Dead vents were frequently observed (e.g., Figure
4m). Temperature adjacent to the active vents was ~12-30ºC with 2.2ºC ambient.
Synaphobranchus eels were constantly present throughout both submersible dives either investigating
active vents, the bait box, or feeding at the bait stations (described above). Heterocarpus shrimp,
benthic ophiuroids, and the chimera H. cf. purpurescens were also observed independent of the bait
stations. Necrophagous lysianassid amphipods (Eurythenes cf. obesus) were not observed (cf. Vetter
& Smith 2005).
Sample Collection & Analyses Biological
Biological and geological samples were collected on each dive (Figure 4, 6). Samples include: 17
coral colonies (Cook = 9, McCall = 8), 6 microbial samples at Lōʻihi (4 microbial mat, 2 adjacent
water column), and 7 rock samples (Cook = 1, McCall = 5, Lōʻihi = 1). Analyses and/or funding
applications for analyses on all sampled material are currently underway. Results thus far suggest that
both Cook and McCall harbour a new species of gorgonian to science; Paramuricea n.sp., fondly
nick-named ‘purple haze’(L. Lamar, Figure 6a).
Discussion
This project provided a rare and unique opportunity to explore two of the oldest seamounts and
adjacent youngest seamount within the Central Pacific. From 43 hours dive (bottom) time >100
species of invertebrates and fishes were observed, and species numbers are expected to increase upon
further investigation. Cook seamount, previously unexplored, yielded a wealth of faunal diversity
similar to that of McCall, including a new species of gorgonian coral, and abundant precious coral
and sponge communities’ characteristic of VMEs (Vulnerable Marine Ecosystems). Fish and
11!
Figure 6. KOK1616 representative biological and geological samples collected. Cook seamount: (a)
Paramuricea n.sp., (b) crusted Mn basalt sample and, (c) Chrysogorgia geniculata attached to rock. McCall
seamount (d) Hemicorallium sp., (e) thinly crusted Mn basalt sample and, (f) Pleurocorallium porcellanum
attached to rock. Lōʻihi seamount (g) Fe-rich venting deposit covered basalt sample, (h) preparing to sample in
a field of chimney vents, (i) sampling microbial mats from beehive vent and, (j) passing over of sample tube
from Pisces IV to V. Images (a, c, d, g-i) M. Cremer, (b, e, f, j) T. Kerby.
12!
scavenger abundance and diversity were surprisingly lower than expected from the visual surveys and
six bait stations conducted. Nevertheless, these observations highlight the necessity to protect these
fragile ecosystems from mining and fishing practices. Exploration at the active submarine volcano
Lōʻihi yielded remarkable fields of finger chimney vents and beehive vents. Part of the vent field
that was previously present on the southeastern wall was obscured, and likely due to a recent
landslide since 2014. It is hoped that results from this brief yet timely expedition will continue to
yield research and conservation dividends beyond those presented here.
Cook & McCall Seamount – Benthic Communities
Current estimates indicate that over 100 species of invertebrates and fishes were observed (Appendix
I). The variety of taxa observed and overall similarities in benthic communities between Cook and
McCall suggest that other factors in addition to gross seamount geomorphology (e.g., conical vs.
ridged) influenced community composition and distribution. For example, observations on the
adjacent Cross and Bishop seamounts reported “dense coral fields” and “high whip coral density” on
the summits “corresponding with high currents” (Kerby T, pers. comm.). Such observations were
~400 m depth and thus occurred at depths above the regional oxygen minimum zone (500 – 1000m;
see Yeh & Drazen 2009). Interestingly, an increased abundance of sponges particularly those within
the Farreidae and Euplectellidae, were present below 930 m depth for both seamounts, which may
also be related to variations in regional ocean chemistry. Further analyses and surveys will determine
depth distribution patterns, with greater abundances reported between 1500-2000 m depth at adjacent
seamounts (Bell et al. 2015).
The benthic communities at Cook and McCall consisted of dense patches of corals (Gorgonians
and/or Scleractinia; Figure 4a) interspersed with sparsely colonised typically unconsolidated terrain
as well as fields of primnoid corals (mainly Calyptrophora wyvillei) on low relief basalt. Gorgonian
octocorals were generally the dominant benthic fauna with some high-density patches of the deep-sea
scleractinian Enallopsammia. Areas of high species abundance and diversity were generally
associated with prominent ridges and wall formations, typically lining the edges and thus likely to
benefit from deep-sea currents. Furthermore, the consistently small colonies of the precious coral
Pleurocorallium porcellanum (Figure 4e) frequently present on the ledges, pinnacles, walls as well as
low relief substratum, may also be indicative of strong current flow through limiting colony height
for this species. Large colonies of Iridogorgia magnispiralis (Figure 4d) and the enigmatic whorls of
the bamboo whip cf. Lepidisis all tended to face eastward with a prevailing current from the west for
example, also suggest unidirectional or a dominant flow regime. Further investigation is necessary,
through deployment of acoustic Doppler current profilers and flow meters, to elucidate the influence
of current dynamics on the ecology and development of keystone benthic marine taxa such as corals
(as discussed by Kelley C., et al. in Bell et al. 2015).
Seamounts have been shown to create localized flow by trapping deep ocean currents, and therefore
recirculating nutrients as well as retaining larvae (Mullineaux & Mills 1997, Roberts et al. 2006).
Such larval retention particularly on the remote seamounts of and around Hawai'i is suggested to lead
to increased endemism (de Forges et al. 2000). For many shallow water and terrestrial groups the
remoteness of the Hawaiian archipelago has indeed led to remarkable endemism (e.g., Kosaki et al.
2016), however, the logistical challenges of deep-sea exploration renders such conclusions difficult
for deep-sea taxa. Whether Paramuricea n.sp., is endemic to the Hawaiian region is unclear.
Certainly, the specimens collected from both Cook and McCall are identical; yet differ considerably
to related material from the Hawaiian archipelago (Rowley SJ, pers. obs.). Increased exploration,
13!
sampling and institutional investigation are necessary to establish this and other Hawaiian deep-sea
taxa as endemic. Similarly, the frequently encountered precious coral P. porcellanum was originally
designated as Corallium kishinouyei by Bayer (1996) collected from the nearby Geologist ‘Cross’
seamount (subsequently placed within the resurrected genus Pleurocorallium by Figueroa & Baco
2014). Having recently been synonymized with P. porcellanum (Tu et al. 2016), and originally
collected from Marcus-Necker Ridge, suggests that this may have a wider distribution than
previously thought (Bayer 1996 also see Ardila et al. 2012). With the exception of highly speciosed
genera such as Narella, evidence for deep-sea endemism for the majority of seamount taxa is
inconclusive and requires further investigation (see Watling et al. 2011). Nevertheless, widely
distributed deep-sea gorgonians and their associate fauna (e.g., Metallogorgia melanotrichos and
Ophiocreas oedipus, Mosher & Watling 2009) were well represented at Cook and McCall.
No evidence of anthropogenic (e.g., destructive fishing practices) disturbance was observed.
However, fish abundance and diversity were lower than expected from the visual surveys and the six
bait stations conducted. This may be due to the patchy benthic communities in many places,
unreported fishing practices, and/or areas and depths surveyed being within the OMZ for the region.
Most notable of the 17 fish taxa observed are the abundance of Synaphobranchus eels, particularly at
the bait stations and throughout the dives at Lōʻihi, the latter likely due to following the bait boxes on
the submersibles, and thus confounding true abundance values! Other notable fish taxa were within
the Ophidiidae and Macrouridae, also recorded at the adjacent Geologist seamounts by Bell et al.
(2015). The Chimera Hydrolagus cf. purpurescens was present on all seamounts, including Lōʻihi.
Both H. cf. purpurescens and the 5-gill Pacific Sleeper shark Somniosus pacificus were observed in
Pele’s Pit, and considered residents over the years (Kerby T., pers. comm.). Of significant note were
the two observations of the charismatic ‘Dumbo’ octopus Grimpoteuthis at Cook, where one was
shedding its skin (desquamation as a mode of excretion through the transfer of melanin; Riley 1994)
although observed phenotypic differences between the two individuals observed imply that they were
not of the same genus, even though are within the suborder Cirrata. More investigation is required to
ascertain the species abundance, diversity and behaviour of this little studied, enigmatic, key predator
group (Villanueva et al. 1997).
Geological history – Fe-Mn crust
This project explored seamounts in unprotected ocean waters, and thus vulnerable to anthropogenic
exploitation. In addition to destructive fishing practices, deep-sea seamounts are of great economic
interest for the mining of mineral-rich (Fe-Mn; Clark et al. 2016) crusts, particularly within the
‘Prime Fe-Mn Crust Zone’ (PCZ). The Fe-Mn crusts of the PCZ are considered one of the most
mineral-rich, often exceeding terrestrial sources by up to 1700 times (e.g., titanium; Hein et al. 2013).
Minerals precipitate from the seawater on to the sediment-free elevated seafloors of numerous
seamounts (Conrad et al. 2016). The large surface area of guyots, such as Cook and McCall, attract
the precipitation of mineral deposits, and are particularly favourable for targeting mineral extraction.
Crustal growth rates can vary between 1-10 mm/m.y (Hein et al. 2000), with a local (Hawaiian)
average ~2.5 mm/m.y (Moore & Clague 2004). The Geologist seamounts have been shown to be of
Late Cretaceous origin (~80 Ma; Sager & Pringle 1987) with Jagger seamount, <10 miles north east
of Cook, and 40 miles north of McCall, having Fe-Mn crusts 80 mm thick (Moore & Clague 2004).
This contrasts with recent evidence from other Geologist seamounts with crusts <2-10 mm thick (Bell
et al. 2015). Crust formation can vary between sample location as well as between the layers within a
single specimen, which may be due to a variety of factors such as depth, hydrodynamic regime,
and/or substratum type (Glasby & Andrews 1977). The variable geomorphology observed at the
14!
Geologist seamounts and interaction with other environmental factors may, therefore, be the source
of such a contrast in Fe-Mn crust thickness. Moreover, such environmental factors may directly or
indirectly explain variations in the benthic community composition observed (also noted by Bell et al.
2015). Whether such intriguing results are due to mass wasting, extremities in flow or sedimentation
relative to geomorphological influence, analytical error or of uncertain geological origin, is unknown
and will remain so unless further exploratory research takes place.
Despite the uncertainty of the age - and therefore geological origin - of these seamounts it is
nonetheless critical to increase our understanding of biological communities associated with Fe-Mn
crusts and the subsequent implications of their loss through industrial mining. Furthermore, valuable
paleoclimatic signals held within the Fe-Mn crust stratigraphy as to the geological origin of these
seamounts will be lost. Here we show that the Geologist seamounts possess distinct geological
features that support a variety of biodiversity, which include new species to science.
Megafaunal Scavengers
The six bait stations conducted throughout this research trip revealed a surprising paucity of
taxonomic diversity. This may most likely be depth related due to decreased oxygen levels for the
region, a pattern also observed by Yeh & Drazen (2009). Other limiting and likely confounding
factors include bait station duration (average 2.43 hr this study cf. average 20.23 hr in the
aforementioned study), location of deployment and/or seasonal pulses in carbon flux and primary
production (but see Drazen 2002). Overnight deployments would most likely reveal greater diversity,
abundance and temporal succession, as well as behaviour including possible scavenger-scavenger
interactions. A notable scavenger-scavenger interaction was the snatching of bait by
Synaphobranchus eels from Heterocarpus shrimp on the summit of McCall seamount. Bait was
snatched prior to the shrimp swimming upwards. Upward swimming was a consistently observed
behaviour from both eels and shrimp once in possession of the prized bait. This suggests that upward
swimming behaviour may be to avoid such competition and/or predation. Interestingly, small eels
were observed swimming against the bait box at an upward angle with bait in mouth for up to 5
minutes before adjusting their angle of swim to actually making way (Rowley, SJ. pers. obs.).
Admittedly the author is not a specialist in this field of research, nevertheless, it would seem that such
swimming behaviour would be a ‘programmed’ response to avoid competition or predation,
compared to that of the sharks observed in this study. High-speed locomotion is energetically
expensive and is generally not selected for in the deep sea, particularly in the absence of light (Seibel
& Drazen 2007). Non-visual predators and scavengers would thus rely on other strategies for
obtaining energy without becoming it (e.g., before the possible onset of a secondary predator
community; Eastman & Thiel 2015). It therefore seems plausible that upward swimming may be a
worthy expenditure of energy to reduce the field of sensory view to other competitors/predators.
Food availability in deep-sea habitats is dependent on primary productivity and large-food fall
(Janßen et al. 2000, Amon et al. 2015). Therefore, scavenger arrival time would be assumed to be
prompt; few scavengers are obligate to a single food source (cf. the amphipod Tryphosa nana
(Krøyer, 1846); Ramsay et al 1997), with fresh carrion of higher nutritional value (Eastman & Thiel
2015). Here, bait stations consisted of diced squid and fish, food items commonly found in the
stomach of large predators and scavengers such as the Pacific sleeper shark, Somniosus pacificus
(Orlov 1999, Yang & Page 1999, Yano et al. 2007), yet this species was not observed to grab or
consume the bait. Similarly, the chimera Hydrolagus cf. purpurescens showed mild interest in the
bait box but did not feed. Only a few observations of the sharks, Etmopterus pusillus actually
15!
consumed the bait. With such observations it lends question to the possible timing (diurnal, seasonal),
type preference or even ‘chase’ of the food for some taxa. Due to their lack of commercial
importance little is known of the life history and distribution of such taxa providing fertile ground
for discovery.
Here, the dominant taxa at all seamounts (Appendix II) were Synaphobranchus eels, and
Heterocarpus shrimp, which were generally present throughout the dives, and thus, may represent
local abundance values as opposed to ‘absence without bait’. Similarly, the smooth lantern sharks E.
pusillus were present at both Cook and McCall bait stations as well as periodically observed
throughout the dives. However, as previously noted the scent of bait from the submersible bait boxes
would confound such observations. The lack of Macrouridae and Ophidiidae may likely be a
consequence of depth, a pattern noted by Janßen et al. (2000) for the former, and only observed at
depths ≥ 1500 m at bait stations conducted within the region (Yeh & Drazen 2009). Occasional
observations throughout the dive surveys however, tends to suggest that population density may not
be the only factor influencing their absence, and thus other factors may be at play such as sensory
ecology, temporal succession or depth-related differences in ocean chemistry. Whether alternative
scavenging communities exist further down the flanks of these seamounts is unknown and certainly
would benefit from further research.
Lōʻihi Revisited Lōʻihi
Lōʻihi seamount, the youngest and most active volcano of the Hawaiian chain, was first documented
in 1940 (Stearns 1946). Macdonald (1952) acknowledged its ‘active’ volcanic status as a
consequence of a large earthquake swarm in 1952. Yet it was not until the latter 1970’s where another
two earthquake swarms motivated an immense research drive until recent years (reviewed by Garcia
et al. 2006). In 1987 HURL conducted the first submersible dives (Pisces V, pilot T. Kerby) initiating
over 20 years pioneering research until 2011. This pioneering geological, geochemical, geophysical
and microbial research shaped the understanding of oceanic island formation to this day.
After more than 5 years since the Pisces’ last visit, and with Lōʻihi still very much active, there was
much question as to what may have changed. A key example and of great research importance, was
the existence and status of Pele’s pit, a ~300 m3 crater formed in the summer of 1996 from an intense
earthquake swarm between July and August (Malahoff 1998, Caplan-Auerbach & Duennebier 2001).
Here it was confirmed that the precarious passage to Pele’s pit still remained well marked and that the
pit itself still remained. However, the previously studied ‘Jet’ site at the southeastern wall of the pit
no longer existed likely due to a recent landslide (i.e., a 2014 expedition, or those prior, do not report
any significant geomorphological changes; Glazer 2014). Where once Pele’s pit was characterised by
200ºC temperatures, active venting and turbid water through suspended volcanic minerals and
bacterial flocks has continued to subside (progressive summaries by Garcia et al. 2006, Glazer &
Rouxel 2009). Active low temperature venting (~12-30ºC with 2.2ºC ambient temperature at present)
increased on ascent up the steep southeastern ridge wall. The observed ‘beehive’ vents leading to
fields of ‘finger’ chimneys demonstrated that volcanic activity was still present with a slight
southeasterly shift at this location.
The morphological variability of microbial mats has been shown to provide microniche stability
relative to oxygen gradients within the actual mats (Chan et al. 2016). Lōʻihi vent fluids are unique
being highly enriched in CO2, CH4, NH4, PO4, Fe, and Mn (reviewed by Garcia et al. 2006), and low
16!
in H2S and O2, the latter due to being situated in the oxygen minimum zone (OMZ; Glazer & Rouxel
2009). This combination of high Fe and CO2, and low H2S and O2, has resulted in a fascinating array
of prokaryotic diversity, with Fe-oxidizing bacteria (FeOB) actually constituting ~60% of the total
Fe-oxidation (Emerson & Moyer 2002). How microbial community composition at beehive vents
differs to the finger chimneys contributing to the overall biological Fe-oxidation at Lōʻihi is
unknown. Greater sampling effort with increased within-mat precision including geochemical factors
such as temperature, O2, CO2, Fe(II) (Glazer & Rouxel 2009, Chan et al. 2016) would further
elucidate how venting communities respond to changes in hydrothermal activity. Of further interest is
how, if at all, such biological Fe-oxidation at Lōʻihi contributes - and perhaps differs to other regions
- to crustal development at the adjacent seamounts is unclear.
Summary
Discovery through deep-sea exploration provides the framework for future research and conservation.
This project reveals evidence of new species discoveries, behavioural interactions, ecological and
geological patterns on little or previously unexplored seamounts. Furthermore changes in
geomorphology, vent type, temperature, and location at Lōʻihi highlight the need for ongoing
geophysical monitoring of hydrothermal developments. Increased precision sampling of the ‘finger’
and ‘beehive’ chimney vents would continue to build on existing microbial knowledge of
biogeochemical pathways and processes which shape the fossil record and our understanding of these
processes over geological time. No evidence of anthropogenic (e.g., destructive fishing practices)
disturbance was observed, yet patchy faunal distributions highlight our lack of understanding of the
abiotic and biotic influences on seamount benthic communities. Similarly, the uncertainty in the
geological origin of the Geologist seamounts would benefit from stratified sampling efforts for
analyses of Fe-Mn crust thickness as well as the assessing valuable paleoclimatic signals stored
within crustal stratigraphic layers. Here, exploration within the ‘Prime Fe-Mn Crust Zone’ (PCZ)
reveals the critical need to research and protect deep-sea seamounts and associate biological
communities from expansive economic demands. By continuously placing mineral resources and
technology above deep-sea biological communities of significant value (e.g., ecosystem function,
pharmaceutical applications), we essentially continue to put ourselves at risk! Thus in a time of
unprecedented yet oxymoronic expansion in green-technology, the unique biological communities of
deep-sea seamounts are vulnerable to extinction. Immediate action is required to protect these
vulnerable marine ecosystems (VMEs), yet only through exploration can substantial evidence furnish
effective protective management decisions.
Acknowledgements
This research project is the result of generous funding of Dr. A. Eustace, Mr. G. Baum of the Baum
Foundation, and logistical support of Conservation International. Sincere appreciation is extended to
the tireless efforts of the captain and crew of RV Ka‘imikai-o-Kanaloa who provided surface support
and exceptional skills in all associated operations. A special thank you to Prof. L. Watling, Prof. S.
Stanley, Prof. S. Donachie, and Prof. C. McFadden for laboratory space, research support and
supplies. We also thank T. Carvalho M.S and Dr. M. Dunlap for SEM support and guidance. Thank
you for the generous donation of the fastCTD profiler from Valeport Ltd, UK. Gratitude is further
extended to Dr. C. Kelley for logistical guidance, and H. Bolick for assistance with accessing
comparative specimen material at the Bernice Pauahi Bishop Museum, USA. Thank you also to N.
Wallsgrove for oven access and Prof. M. Garcia for geological sample guidance, Dr. R. Pyle for
comments on the report, and Dr. D. Ebert for Etmopterus pusillus identification. Research was
conducted outside state waters, and thus permits were not required.
17!
Abbreviations & Glossary
AIVL - Advanced Imaging & Visualization Laboratory (of WHOI), USA
AP – Associated Press, USA
BF – Baum Foundation, USA
CI – Conservation International
GIS – Geographic Information System
HURL – Hawai‘i Undersea Research Laboratory, USA
UH – University of Hawai‘i at Manoa, USA
WHOI – Woods Hole Oceanographic Institution, USA
References
Amon DJ, Copley JT, Dahlgren TG, Horton T, Kemp KM, Rogers AD, Glover AG. 2015. Observations of
fauna attending wood and bone deployments from two seamounts on the Southwest Indian Ridge. Deep-Sea
Res Pt II.!http://dx.doi.org/10.1016/j.dsr2.2015.07.003i
Ardila N, Giribet G, Sanchez J. 2012. A time-calibrated molecular phylogeny of the precious corals:
reconciling discrepancies in the taxonomic classification and insights into their evolutionary history. BMC
Evol Biol. 12: 246.
Bayer FM. 1996. Three new species of precious coral (Anthozoa: Gorgonacea, genus Corallium) from Pacific
waters. Proc Biol Soc Wash. 109: 205–28.
Bell KLC, Brennan ML, Flanders J, Raineault NA, Wagner K, eds. 2016. New frontiers in ocean exploration:
The E/V Nautilus and NOAA Ship Okeanos Explorer 2015 field season. Oceanography. 29(1),
supplement, 84: http://dx.doi.org/10.5670/oceanog.2016.supplement.01.
Caplan-Auerbach J, Duennebier F. 2001. Seismicity and velocity structure of Lōʻihi seamount from the 1996
earthquake swarm. Bull Seis Soc Am. 91(2): 178-190.
Chan CS, McAllister SM, Leavitt AH, Glazer BT, Krepski ST, Emerson D. 2016. The architecture of iron
microbial mats reflects the adaptation of chemolithotrophic iron oxidation in freshwater and marine
environments. Front Microbiol. 7:796. doi: 10.3389/fmicb.2016.00796
Clark MR, Althaus F, Schlacher TA, Williams A, Bowden DA, Rowden AA. 2016. The impacts of deep-sea
fisheries on benthic communities: a review. ICES Journal of Marine Science. 73: i51–i69.
Conrad T, Hein JR, Paytan A, Clague DA. 2016. Formation of Fe-Mn crusts within a continental margin
environment. Ore Geol. Rev. http://dx.doi.org/10.1016/j.oregeorev.2016.09.010
de Forges BR, Koslow JA, Poore GC. 2000. Diversity and endemism of the benthic seamount fauna in the
southwest Pacific. Nature. 405(6789): 944-7.
Donachie SP, Hou S, Lee KS, Riley CW, Pikina A, Belisle C, Kempe S, Gregory TS, Bossuyt A, Boerema J,
Liu J. 2004. The Hawaiian Archipelago: a microbial diversity hotspot. Microbial Ecol. 48(4): 509-520.
Drazen JC. 2002. A seasonal analysis of the nutritional condition of deep-sea macrourid fishes in the north-
east Pacific. J. Fish Biol. 60: 1280-1295.
18!
Dymond J, Windom HL. 1968. Cretaceous K-Ar ages from Pacific Ocean seamounts. Earth Planet Sc Lett.
4(1): 47-52.
Eakins BW, Robinson JE, Kanamatsu T, Naka J, Smith JR, Takahashi E, Clague DA. 2003, Hawai‘i's
volcanoes revealed: U.S. Geological Survey Geologic Investigations Series Map I-2809, 1 plate,
https://pubs.usgs.gov/imap/2809/.
Eastman LB, Thiel M. 2015. Foraging behaviour of crustacean predators and scavengers. In M Thiel, L
Watling (Ed.), Lifestyles and Feeding Biology: Volume II. (pp. 535-556) Oxford University Press, USA.
Emerson D, Moyer CL. 2002. Neutrophilic Fe-oxidizing bacteria are abundant at the Lō’ihi Seamount
hydrothermal vents and play a major role in Fe oxide deposition. App Environ Microbiol. 68(6): 3085-93.
Emerson D, Scott JJ, Leavitt A, Fleming E, Moyer C. 2017. In situ estimates of iron-oxidation and accretion
rates for iron-oxidizing bacterial mats at Lō’ihi Seamount. bioRXiv doi: https://doi.org/10.1101/095414
Figueroa DF, Baco AR. 2014. Complete mitochondrial genomes elucidate phylogenetic relationships of the
deep-sea octocoral families Coralliidae and Paragorgiidae. Deep Sea Res Part II. 99: 83–91.
Garcia MO, Caplan-Auerbach J, De Carlo EH, Kurz MD, Becker N. 2006. Geology, geochemistry and
earthquake history of Lōʻihi seamount, Hawai‘i’s youngest volcano: Chemie der Erde. 66: p. 81–108,
doi: 10 .1016 /j .chemer .2005 .09 .002 .
Garcia MO, Smith JR, Tree JP, Weis D, Harrison L, Jicha BR. 2015. Petrology, geochemistry, and ages of
lavas from Northwest Hawaiian Ridge volcanoes, in Neal CR, Sager WW, Sano T, Erba E. eds., The Origin,
Evolution, and Environmental Impact of Oceanic Large Igneous Provinces: Geological Society of America
Special Paper. 511: 1–25.
Glasby GP, Andrews JE. 1977. Manganese crusts and nodules from the Hawaiian Ridge: Pacific Science: 31:
363–379.
Glazer BT. 2014. Hydrothermal Iron Biogeochemistry and Microbial Habitats at Lōʻihi Seamount. R/V
Falkor/AUV Sentry Expedition FK140626 Cruise Report. 1-137
Glazer BT, Rouxel OJ. 2009. Redox speciation and distribution within diverse iron-dominated microbial
habitats at Lōʻihi Seamount. Geomicrobiol J. 26(8): 606-622.
Hein JR, Conrad TA, Dunham RE. 2009. Seamount characteristics and mine-site model applied to exploration-
and mining-lease-block selection for cobalt-rich ferromanganese crusts. Mar Geores Geotech. 27(2): 160-76.
Hein JR, Mizell K, Koschinsky A, Conrad TA. 2013. Deep-ocean mineral deposits as a source of critical
metals for high- and green-technology applications: Comparison with land-based resources. Ore Geol Rev. 51:
1-14.
Janßen F, Treude T, Witte U. 2000. Scavenger assemblages under different trophic conditions: a case study in
the deep Arabian Sea. Deep-Sea Res. 47: 2999-3026.
Karl DM, Brittain AM, Tilbrook BD. 1989. Hydrothermal and microbial processes at Lōʻihi Seamount, a mid-
plate hot-spot volcano. Deep-Sea Res. 36(11): 1655-1673.
19!
Kosaki RK, Pyle RL, Leonard JC, Hauk BB, Whitton RK, Wagner D. 2016. 100% endemism in mesophotic
reef fish assemblages at Kure Atoll, Hawaiian Islands. Mar Biodiv. 1-2.
Last PR, Stevens JD. 1994. Sharks and Rays of Australia. Melbourne. CSIRO Division of Fisheries. 513p.
Leray M, Knowlton N. 2016. Censusing marine eukaryotic diversity in the twenty-first century. Phil. Trans. R.
Soc. B. 371: 20150331.
Macdonald GA. 1952. The south Hawai‘i earthquakes of March and April, 1952: Volcano Letter. 515: 1–5.
Malahoff A. 1998. An integrated shipboard ocean floor research system using a SeaBeam 210 ocean
survey system installed on the R/V Ka’imikai-o-Kanaloa. Mar Geodesy. 21: 97–109.
McFadden CS, Benayahu Y, Pante E, Thoma JN, Nevarez PA, France SC. 2011. Limitations of mitochondrial
gene barcoding in Octocorallia. Mol Ecol Res. 11:19–31.
McFadden CS, Brown AS, Brayton C, Hunt CB, van Ofwegen LP. 2014. Application of DNA barcoding in
biodiversity studies of shallow-water octocorals: molecular proxies agree with morphological estimates of
species richness in Palau. Coral Reefs. 33: 275-286.
Moore JG, Clague DA. 2004. Hawaiian submarine manganese-iron oxide crusts—A dating tool?
Geol Soc
Am
Bull.
116(3-4): 337-47.
Mosher C, Watling L. 2009. Partners for life: A brittle star and its octocoral host. Mar Ecol Prog Ser. 397: 81-
88.
Mullineau LS, Mills SW. 1997. A test of the larval retention hypothesis in seamount-generated flows. Deep-
Sea Res PT I. 44(5): 745-70.
Muzik KM. 1979. Systematic revision of the Hawaiian Paramuriceidae and Plexauridae. Thesis, University of
Miami, Coral Gables, Florida, USA, pp 227.
Orlov AM. 1999. Capture of especially large sleeper shark Somniosus pacificus (Squalidae) with some notes
on its ecology in northwestern Pacific. J Ichthyol. 39: 548–553.
R. 2013. A language environment for statistical computing.
Ramsay K, Kaiser MJ, Moore PG, Hughes RN. 1997. Consumption of fisheries discards by benthic
scavengers: utilization of energy subsidies in different marine habitats. J.Anim Ecol. 884-96.
Riley PA. 1994. Photopigmentation. In Photobiology in Medicine. Springer US. pp. 99-112.
Roberts JM, Wheeler AJ, Freiwald A. 2006. Reefs of the deep: the biology and geology of cold-water coral
ecosystems. Science. 312(5773): 543-7.
Sager WW, Pringle MS. 1987. Paleomagnetic constraints on the origin and evolution of the Musicians
and South Hawaiian seamounts, central Pacific Ocean. 133–162 in
Seamounts, Islands, and Atolls.
AGU Geophysical Monograph Series, vol. 43, Keating B, Fryer P, Batiza R, Boehlert G, eds, American
Geophysical Union, Washington, DC.
20!
Seibel BA, Drazen JC. 2007. The rate of metabolism in marine animals: environmental constraints, ecological
demands and energetic opportunities. Phil. Trans. R. Soc. B. 362(1487): 2061-78.
Stearns HT. 1946. Geology of the Hawaiian Islands. Hawai‘i. Div Hydrogr Bull. 8, 112 p
Tu TH, Dai CF, Jeng MS. 2016. Taxonomic revision of Coralliidae with descriptions of new species from New
Caledonia and the Hawaiian Archipelago. Mar Biol Res. 12:10, 1003-1038, DOI:
10.1080/17451000.2016.1241411
Villanueva R, Segonzac M, Guerra A. 1997. Locomotion modes of deep-sea cirrate octopods (Cephalopoda)
based on observations from video recordings on the Mid-Atlantic Ridge. Mar Biol. 129(1): 113-22.
Watling L, France SC, Pante E, Simpson A. 2011. Biology of deep-water octocorals. Adv Mar Biol.
60:41-122.
Yang M, Page BN. 1999. Diet of Pacific sleeper shark, Somniosus pacificus, in the Gulf of Alaska.
Fish B-
NOAA
. 97: 406–409.
Yano K, Stevens JD, Compagno LJ. 2007. Distribution, reproduction and feeding of the Greenland
shark Somniosus (Somniosus) microcephalus, with notes on two other sleeper sharks, Somniosus
(Somniosus) pacificus and Somniosus (Somniosus) antarcticus. J Fish Biol. 70(2): 374-90.
Yeh J, Drazen JC. 2009. Depth zonation and bathymetric trends of deep-sea megafaunal scavengers of the
Hawaiian Islands. Deep-Sea Res. 56: 251-266.
Yu L, Zhang W, Liu L, Yang J. 2015. Determining Microeukaryotic Plankton Community around Xiamen
Island, Southeast China, Using Illumina MiSeq and PCR-DGGE Techniques. PLoS ONE.
10(5): e0127721. doi:10.1371/journal.pone.0127721
21!
Appendix I. KOK1616 expedition preliminary species survey observation table. Relative species abundance was estimated using ACFOR
where A = Abundance > 50 individuals observed in a dive; C = Common ≤ 50 and >20 individuals observed; F = Frequent ≤ 20 and >10
individuals observed; O = Occasional ≤ 10 and >2 individuals observed; R= Rare ≤ 2 individuals observed. OTU, Operational Taxonomic
Unit. Indet., indeterminate. Taxon identification is an approximation when not sampled, and thus should be treated with caution.
Phylum
Class
Subclass
OTU
Seamount
Order
Family
Genus
species
Cook
McCall
Lōʻihi
Annelida
Polychaeta
Indet.
-
-
-
-
-
O
-
Annelida
Polychaeta
Sedentaria
Sabellida
Serpulidae
Indet.
-
-
R
-
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Indet.
-
-
O
O
-
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Acanthephyridae
Acanthephyra
sp.
O
R
R
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Aristeidae
Plesiopenaeus cf.
sp.
F
R
-
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Chirostylidae
Uroptychus
sp.
C
O
-
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Eumunididae
Indet.
-
R
R
-
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Galatheidae
-
-
A
C
-
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Homolidae
-
-
-
R
-
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Munididae
-
-
F
R
-
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Nematocarcinidae
Nematocarcinus
tenuirostris
F
C
F
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Pandalidae
Heterocarpus
sp.
O
O
F
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Pandalidae
Heterocarpus
ensifer cf.
F
F
O
Arthropoda
Malacostraca
Eumalacostraca
Decapoda
Pandalidae
Heterocarpus
laevigatus
F
F
O
Chordata
Thaliacea
-
Salpa
Salpidae
Indet.
-
-
O
-
Chordata
Actinopteri
-
Anguilliformes
Indet.
-
-
-
O
R
Chordata
Actinopteri
-
Anguilliformes
Synaphobranchidae
Indet.
-
-
O
R
Chordata
Actinopteri
-
Anguilliformes
Synaphobranchidae
Synaphobranchus
sp.
C
C
C
Chordata
Actinopteri
-
Anguilliformes
Synaphobranchidae
Synaphobranchus
brevidorsalis
O
O
O
Chordata
Actinopteri
-
Gadiformes
Macrouridae
-
-
O
O
-
Chordata
Actinopteri
-
Gadiformes
Macrouridae
Gadomus cf.
sp.
F
-
-
Chordata
Actinopteri
-
Gadiformes
Macrouridae
Coryphaenoides
sp.
F
-
-
Chordata
Actinopteri
-
Gadiformes
Macrouridae
Nezumia
sp.
O
-
-
Chordata
Actinopteri
-
Gadiformes
Moridae
Guttigadus
sp.
R
-
-
Chordata
Actinopteri
-
Lophiiformes
Lophiidae
Sladenia
remiger
R
-
-
Chordata
Actinopteri
-
Ophidiiformes
Ophidiidae
-
-
O
-
R
Chordata
Actinopteri
-
Ophidiiformes
Ophidiidae
Lamprogrammus
sp.
O
O
-
Chordata
Actinopteri
-
Ophidiiformes
Ophidiidae
Lamprogrammus
brunswigi
R
-
-
Chordata
Holocephali
-
Chimaeriformes
Chimaeridae
Indet.
-
-
R
-
Chordata
Holocephali
-
Chimaeriformes
Chimaeridae
Hydrolagus
purpurescens
R
R
R
22!
Phylum
Class
Subclass
OTU
Seamount
Order
Family
Genus
species
Cook
McCall
Lōʻihi
Chordata
Elasmobranchii
Neoselachii
Squaliformes
Etmopterus pusillus
O
C
-
Chordata
Elasmobranchii
Neoselachii
Squaliformes
Somniosus
pacificus
-
-
R
Cnidaria
Anthozoa
Hexacorallia
Actinaria
sp.
-
O
-
Cnidaria
Anthozoa
Hexacorallia
Antipatharia
sp.
O
O
-
Cnidaria
Anthozoa
Hexacorallia
Scleractinia
-
F
-
-
Cnidaria
Anthozoa
Hexacorallia
Scleractinia
-
Bathypathes
Indet.
Enallopsammia
sp.
A
C
-
Cnidaria
Anthozoa
Hexacorallia
Zoantharia
-
-
O
O
-
Cnidaria
Anthozoa
Hexacorallia
Zoantharia
-
-
C
O
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
-
-
F
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Anthmastus
sp.
-
R
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Acanthogorgia
sp.
C
F
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Corallium
sp.
F
F
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Hemicorallium
sp.
F
F
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Pleurocorallium
porcellanum
A
A
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Chrysogorgia
sp.
C
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Chrysogorgia
geniculata
F
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Iridogorgia
sp.
R
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Iridogorgia
bella
F
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Iridogorgia
magnispiralis
C
O
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Iridogorgia
splendens
C
O
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Etmopteridae
Somniosidae
Exocoelactinidae
Schizopathidae
Dendrophylliidae
Dendrophylliidae
- Parazoanthidae
Indet.
Alcyoniidae
Acanthogorgiida
e Coralliidae
Coralliidae
Coralliidae
Chrysogorgiidae
Chrysogorgiidae
Chrysogorgiidae
Chrysogorgiidae
Chrysogorgiidae
Chrysogorgiidae
Chrysogorgiidae
Metallogorgia
melanotrichos
F
O
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Isididae
-
-
C
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Isididae
Acanella
sp.
F
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Isididae
Lepidisis
sp.
C
O
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Paragorgiidae
Paragorgia
sp.
R
R
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Plexauridae
-
-
O
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Plexauridae
Paramuricea
n.sp.
C
C
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Primnoidae
-
-
-
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Primnoidae
Callogorgia
cf. gilberti
O
R
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Primnoidae
Calyptrophora
sp.
R
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Primnoidae
Calyptrophora
wyvillei
A
F
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Primnoidae
Candidella
gigantea cf.
F
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Primnoidae
Narella
sp.
C
F
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Primnoidae
Narella
dichotoma cf.
F
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Primnoidae
Narella
gigas cf.
O
-
-
23!
Phylum
Class
Subclass
OTU
Seamount
Order
Family
Genus
species
Cook
McCall
Lōʻihi
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Primnoidae
Narella
macrocalyx
O
-
-
Cnidaria
Anthozoa
Octocorallia
Alcyonacea
Primnoidae
Paracalyptrophora
sp.
O
R
-
Cnidaria
Anthozoa
Octocorallia
Pennatulacea
-
-
-
O
R
-
Cnidaria
Hydrozoa
Hydroidolina
Anthoathecata
Stylasteridae
-
-
R
R
-
Cnidaria
Hydrozoa
Trachylinae
Trachymedusae
Rhopalonematidae
Crossota
sp.
O
O
-
Cnidaria
Scyphozoa
Indet.
-
-
-
-
-
-
R
Ctenophora
Tentaculata
-
Lobata
Indet.
-
-
-
R
R
Ctenophora
Tentaculata
-
Lobata
Bathocyroidae
Bathocyroe
sp.
O
F
R
Echinodermata
Asteroidea
-
Spinulosida
Echinasteridae
Indet.
-
R
F
-
Echinodermata
Asteroidea
-
Valvatida
Goniasteridae
Indet.
-
F
O
-
Echinodermata
Asteroidea
-
Valvatida
Goniasteridae
Calliaster
sp.
O
-
-
Echinodermata
Crinoidea
-
-
-
Echinodermata
Echinoidea
Euechinoidea
Echinothurioida
Indet.
-
-
O
R
-
Echinodermata
Holothuroidea
-
Aspidochirotida
Synallactidae
Hansenothuria
sp.
-
O
-
Echinodermata
Ophiuroidea
-
Ophiurida
Ophiuridae
-
-
F
-
O
Echinodermata
Ophiuroidea
-
Ophiurida
Ophiacanthidae
-
-
C
-
-
Echinodermata
Ophiuroidea
-
Euryalida
Asteroschematidae
-
-
C
C
-
Echinodermata
Ophiuroidea
-
Euryalida
Asteroschematidae
Asteroschema
sp.
A
C
-
Echinodermata
Ophiuroidea
-
Euryalida
Asteroschematidae
Ophiocreas
sp.
O
O
-
Echinodermata
Ophiuroidea
-
Euryalida
Asteroschematidae
Ophiocreas
oedipus
F
-
-
Mollusca
Bivalva
Indet.
-
-
-
-
-
R
-
Mollusca
Cephalopoda
Coleoidea
Octopoda
Opisthoteuthidae
Grimpoteuthis
sp.
R
-
-
Porifera
Indet.
-
-
-
-
-
C
O
-
Porifera
Hexactinellida
Amphidiscophora
Amphidiscosida
Pheronematidae
-
-
F
R
-
Porifera
Hexactinellida
Amphidiscophora
Amphidiscosida
Pheronematidae
Semperella cf.
sp.
-
R
-
Porifera
Hexactinellida
Amphidiscophora
Amphidiscosida
Pheronematidae
Poliopogon cf.
sp.
-
O
-
Porifera
Hexactinellida
Hexasterophora
Indet.
-
-
-
F
O
-
Porifera
Hexactinellida
Hexasterophora
Hexactinosida
Auloplacidae
Auloplax cf.
sp.
R
O
-
Porifera
Hexactinellida
Hexasterophora
Hexactinosida
Euretidae
-
-
C
O
-
Porifera
Hexactinellida
Hexasterophora
Hexactinosida
Euretidae
Eurete
sp.
O
O
-
Porifera
Hexactinellida
Hexasterophora
Hexactinosida
Farreidae
Indet
-
F
C
-
Porifera
Hexactinellida
Hexasterophora
Hexactinosida
Farreidae
Aspidoscopulia cf.
sp.
R
R
-
24!
Phylum
Class
OTU
Seamount
Subclass
Order
Family
Genus
species
Cook
McCall
Lōʻihi
Porifera
Hexactinellida
Hexasterophora
Hexactinosida
Farreidae
Farrea
sp.
R
F
-
Porifera
Hexactinellida
Hexasterophora
Lyssacinosida
Indet.
-
-
-
R
-
Porifera
Hexactinellida
Hexasterophora
Lyssacinosida
Euplectellidae
-
-
F
R
-
Porifera
Hexactinellida
Hexasterophora
Lyssacinosida
Euplectellidae
Bolosoma
sp.
-
R
-
Porifera
Hexactinellida
Hexasterophora
Lyssacinosida
Euplectellidae
Corbitella
sp.
C
C
-
Porifera
Hexactinellida
Hexasterophora
Lyssacinosida
Euplectellidae
Dictyaulus
starmeri
R
O
-
Porifera
Hexactinellida
Hexasterophora
Lyssacinosida
Euplectellidae
Euplectella
sp.
R
O
-
Porifera
Hexactinellida
Hexasterophora
Lyssacinosida
Euplectellidae
Regadrella
sp.
-
R
-
Porifera
Hexactinellida
Hexasterophora
Lyssacinosida
Euplectellidae
Walteria
flemmingi
-
O
-
Porifera
Hexactinellida
Hexasterophora
Lyssacinosida
Rossellidae
Caulophacus cf.
sp.
-
R
-
Porifera
Hexactinellida
Hexasterophora
Aulocalycoida
Uncinateridae
Tretopleura
sp.
F
C
-
25!
Appendix II. KOK1616 summary table of bait station fauna present and actively scavenging (bait handled and/or actively consumed).
OTU, Operational Taxonomic Unit; n, number of individuals present; scav., number of individuals actively scavenging.
OTU
Cook
McCall
Lōʻihi
Order
Family
Genus
species
1007 m
1053 m
875 m
1156 m
1324 m
1324 m
n
Scav.
n
Scav.
n
Scav.
n
Scav.
n
Scav.
n
Scav.
Decapoda
Nematocarcinus
tenuirostris
-
-
2
2
1
1
-
-
-
-
-
-
Decapoda
Heterocarpus
sp.
-
-
-
-
-
-
-
-
-
-
4
4
Decapoda
Heterocarpus
ensifer cf.
4
4
1
1
-
-
1
1
3
3
-
-
Decapoda
Heterocarpus
laevigatus
-
-
-
-
4
4
4
4
1
1
-
-
Squaliformes
Etmopterus pusillus
5
4
5
4
3
0§
5
2§
-
-
-
-
Squaliformes
Nematocarcinidae
Pandalidae
Pandalidae
Pandalidae
Etmopteridae
Somniosidae
Somniosus
pacificus
-
-
-
-
-
-
-
-
1*
-
1*
-
Chimaeriformes
Chimaeridae
Hydrolagus
purpurescens
-
-
-
-
1
-
-
-
-
-
-
-
Anguilliformes
Synaphobranchidae
Synaphobranchus
sp.
3
3
3
3
7
7
0§
0§
13‡
13‡
6
6
*Depicts the same individual observed at two different bait stations.
§Submersible camera not fully aimed on the entire bait station, data gleaned from what could be observed and pilot/observer comments.
‡10 juveniles (≤ 25 cm) observed
26!
Appendix III. KOK1616 Biological sample inventory for coral (Octocorallia and Hexacorallia) collected at Cook and McCall Seamounts,
and microbial communities at Lōʻihi. Sample code: P4, Pisces IV; P5, Pisces V; submersible dive number. Preservation: 95% ethanol; Z-Fix,
ionized zinc-formalin fixative; DMSO, Dimethyl sulfoxide; RNAlater, RNA stabilization and storage. Phylogenetic markers: mutS,
mitochondrial marker; 28S & ITS, nuclear markers; 16S, ribosomal marker used for Prokaryotes; 18S, ribosomal marker used for
Eukaryotes.
Sample #
Depth
(m)
Location
Taxonomy
Preservation
Marker(s)
Latitude
Longitude
Family
Genus
species
95%
Z-Fix
DMSO
RNAlater
Cook Seamount - 06.09.2016
P5-859 Spec.4
1013
N19:17.891
W157:09.915
Dendrophylliidae
Enallopsammia
sp.
Y
Y
Y
Y
-
P4-298 Spec.1
935
N18:43.032
W157:04.880
Plexauridae
Paramuricea
Y
Y
Y
Y
16S/18S/ITS
P4-298 Spec.2
1020
N19:18.098
W157:09.977
Primnoidae
Calyptrophora
Y
Y
Y
Y
mutS/28S
P4-298 Spec.3
1062
N19:18.002
W157:09.913
Chrysogorgiidae
Chrysogorgia
Y
Y
Y
Y
mutS/28S
P4-298 Spec.4
1071
N19:17.996
W157:09.886
Isididae
Lepidisis
Y
Y
Y
Y
-
P4-298 Spec.5
1129
N19:17.721
W157:09.769
Acanthogorgiidae
Acanthogorgia
n.sp.
wyvillei
geniculata
sp.
sp.
Y
Y
Y
Y
-
P5-859 Spec.1
1014
N19:18.077
W157:10.031
Plexauridae
Paramuricea
Y
Y
Y
Y
mutS/28S
16S/18S/ITS
P5-859 Spec.2
1022
N19:17.908
W157:09.947
Primnoidae
Callogorgia
Y
Y
Y
Y
mutS/28S
P5-859 Spec.3
1027
N19:17.922
W157:09.921
Primnoidae
Calyptrophora
n.sp.
gilberti
wyvillei
Y
Y
Y
Y
mutS/28S
McCall Seamount - 07.09.2016
P4-299 Spec.7
932
N18 43.070
W157 04.647
Dendrophylliidae
Enallopsammia
sp.
Y
Y
Y
Y
-
P5-860 Spec.2
923
N18 42.995
W157 04.871
Dendrophylliidae
-
sp.
Y
Y
Y
Y
-
P4-299 Spec.1
935
N18:43.032
W157:04.880
Coralliidae
Hemicorallium
sp.
Y
Y
Y
Y
-
P4-299 Spec.2
911
N18:42.983
W157:04.815
Dendrophylliidae
Enallopsammia
sp.
Y
Y
Y
Y
-
P4-299 Spec.3
878
N18:42.989
W157:04.765
Plexauridae
Paramuricea
n.sp.
Y
Y
Y
Y
-
P4-299 Spec.4
878
N18:42.989
W157:04.765
Coralliidae
Pleurocorallium
porcellanum
Y
Y
Y
Y
mutS/28S
P4-299 Spec.6
932
N18 43.070
W157 04.647
Dendrophylliidae
Enallopsammia
sp.
Y
Y
Y
Y
-
P5-860 Spec.1
875
N18:42.912
W157:04.762
Coralliidae
Pleurocorallium
porcellanum
Y
Y
Y
Y
-
Lōʻihi Seamount - 07.09.2016
Cultured Media
P5-861 HIB
1291
N18 54.345
W155 15.402
Adjacent water
Y
-
-
Y
Y
16S/18S/ITS
P5-861 H0
1285
N18 54.389
W155 15.398
Adjacent water
Y
-
-
Y
Y
16S/18S/ITS
P5-861 M35
1285
N18 54.345
W155 15.402
Microbial mat (Finger chimney vent)
Y
-
-
Y
Y
16S/18S/ITS
P5-861 M43
1285
N18 54.345
W155 15.402
Microbial mat (Finger chimney vent)
Y
-
-
Y
Y
16S/18S/ITS
P5-861 M53
1295
N18 54.389
W155 15.398
Microbial mat (Beehive vent)
Y
-
-
Y
Y
16S/18S/ITS
P5-861 M05
1295
N18 54.389
W155 15.398
Microbial mat (Beehive vent)
Y
-
-
Y
Y
16S/18S/ITS
27!
Appendix IV. KOK1616 geological sample inventory collected at the seamounts Cook, McCall
and Lōʻihi. Sample code: P4, Pisces IV; P5, Pisces V; submersible dive number. Asterisk denotes
samples used for further analyses at the University of Hawai‘i at Manoa, USA (see Garcia et al.
2015).
Sample #
Depth (m)
Location
Sample Type
Preservation
Latitude
Longitude
Dried 60°C
Cook Seamount - 06.09.2016
P5-859 Rock 1*
1003
N19:18.112
W157:09.963
Mn-crusted Basalt
Y
McCall Seamount - 07.09.2016
P4-299 Rock1*
911
N18:42.983
W157:04.815
Thinly Mn-crusted Basalt
Y
P4-299 Rock2*
911
N18:42.983
W157:04.815
Thinly Mn-crusted Basalt
Y
P4-299 Rock3*
932
N18 43.070
W157 04.647
Thinly Mn-crusted Basalt
Y
P5-860 Rock1
1003
N18:43.081
W157:04.963
Thinly Mn-crusted Basalt
Y
P5-860 Rock2
1028
N18 43.301
W157 04.841
Thinly Mn-crusted Basalt
Y
Lōʻihi Seamount - 08.09.2016
P4-300 Rock1*
1158
N18:54.360
W155:15.777
Fe-rich vent deposit
covered basalt
Y
